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A systematic revision of the South African Pettalidae(Arachnida Opiliones Cyphophthalmi) based on a combinedanalysis of discrete and continuous morphological characterswith the description of seven new species
Benjamin L de BivortABC and Gonzalo GiribetB
ARowland Institute at Harvard Harvard University 100 Edwin Land Boulevard Cambridge MA 02142 USABMuseum of Comparative Zoology and Department of Organismic and Evolutionary BiologyHarvard University 26 Oxford Street Cambridge MA 02138 USA
CCorresponding author Email debivortrowlandharvardedu
Abstract The cyphophthalmid family Pettalidae in South Africa is revised and seven new species are described frommuseummaterial collected between 1939 and 1985 Two of these are placed in the genusPurcellia andfive inParapurcelliabringing the total number of described South African cyphophthalmids to 15 In addition Purcellia peregrinator istransferred to the genus Parapurcellia Phylogenetic analyses of discrete morphological and continuous morphometriccharacters both separately and in combination support the generic assignments and contribute towards a more detailedunderstanding of the systematics of the group in South Africa In order to assess the stability of our phylogenetic results thedifferent morphological datasets were analysed under equal and implied weighting as well as under several weightingschemes that varied the respective contribution to tree length of the discrete and continuous data partitions These variationsgenerated two phylogenetic hypotheses (1) monophyly of the South African pettalids +Austropurcellia from north-easternAustralia as a derived clade within Pettalidae and (2) polyphyly of the South African pettalids with Parapurcellia basalwithin Pettalidae The latter hypothesis is congruent with previous molecular phylogenies of Cyphophthalmi and hasmoderate bootstrap support The sisterhood of Purcellia griswoldi sp nov and P lawrencei sp nov receives high nodalsupport across analytic methods New combination Parapurcellia peregrinator (Lawrence 1963)
Additional keywords Afromontane forests Gondwanan biogeography morphometrics South Africa
Introduction
The opilionid order Cyphophthalmi (often referred to as miteharvestmen) is represented by 17 species in Africa (Giribet2000 Giribet and Prieto 2003) These include members inthe tropical families Neogoveidae (three described species inthe genus Parogovia Hansen 1921) represented in westernequatorial and western sub-Saharan Africa (Sierra LeoneIvory Coast Cameroon Equatorial Guinea and GabonHansen 1921 Juberthie 1969 Legg 1990 Giribet 2000authorsrsquo unpubl data) and Ogoveidae (three described speciesin the genus Ogovea Roewer 1923) represented in westernequatorial Africa (Cameroon Equatorial Guinea and GabonHansen and Soslashrensen 1904 Hansen 1921 Giribet 2000Giribet and Prieto 2003) plus a species of uncertain affinityfrom KenyaMarwe coarctata Shear 1985 (Shear 1985 Giribetand Boyer 2002) Within the territories of the former temperateGondwana the family Pettalidae is represented in South Africaby three genera the monotypic Speleosiro Lawrence 1931 andseven species in the genera Purcellia Hansen amp Soslashrensen 1904and Parapurcellia Rosas Costa 1950 plus a species in the
monotypic genus Manangotria Shear amp Gruber 1996 inMadagascar A second monotypic genus Ankaratra Shear ampGruber 1996 remains untested phylogenetically and is so farconsidered incertae sedis (Shear andGruber 1996 Giribet 2000)
Pettalidae contains eleven genera distributed across thecontinental fragments of temperate Gondwana (Boyer andGiribet 2007 Giribet and Boyer 2007) Chileogovea Roewer1961 (Chile) ndashAorakiBoyerampGiribet 2007RakaiaHirst 1925andNeopurcellia Forster 1948 (NewZealand) ndashAustropurcelliaJuberthie 1988 (QueenslandAustralia) ndashKarripurcelliaGiribet2003 (Western Australia) ndash Pettalus Thorell 1876 (Sri Lanka) ndashand the abovementionedAfrican genera Each genus is restrictedto a single landmass despite prior hypotheses of trans-Tasmandistributions in Rakaia and Neopurcellia reported from NewZealand and Australia (see Boyer and Giribet 2007)
There is considerable molecular and morphological supportfor the monophyly of many family-level clades within theCyphophthalmi (Giribet and Boyer 2002 Boyer et al 2007a)but the intergeneric relationships especially within the familyPettalidae are less resolved (eg Giribet 2003 Boyer and Giribet
CSIRO 13 December 2010 101071IS10015 1445-522610040371
CSIRO PUBLISHING
wwwpublishcsiroaujournalsis Invertebrate Systematics 2010 24 371ndash406
2007 2009 de Bivort et al 2010) Molecular evidence favoursthe monophyly of all genera as currently defined although only afew (Aoraki Rakaia Pettalus) have been thoroughly sampled inmolecular studies (Boyer and Giribet 2007 Boyer et al 2007a2007b) In these studies the African pettalids were restricted torepresentatives of only three species one in the genus Purcelliaand two in the genus Parapurcellia and South Africa wasparaphyletic with respect to other landmasses
The described South African pettalid fauna is currentlydistributed in two geographic regions (Fig 1) Two speciesare found in the Western Cape province specifically in andaround Table Mountain Purcellia illustrans Hansen ampSoslashrensen 1904 has been collected in forests surroundingTable Mountain Speleosiro argasiformis Lawrence 1931 isa large (5mm) cyphophthalmid found in the Wynberg cavesystem in Table Mountain (Juberthie 1970) The formerspecies is a medium-sized cyphophthalmid with morphologicalcharacteristics typical of other members of South Africanpettalids such as relatively unmodified opisthosomal posteriortergites in males and females second palp articles that are lessthan half ornamented and bilobed anal plates in males Forillustrations of these and other cyphophthalmid morphologicalterms see de Bivort et al (2010) One conspicuous featureof P illustrans is the position of its anal plate which is shiftedanteriorly onto the ventral surface of the animal comparedwith other pettalids Speleosiro argasiformis has conspicuoustroglomorphic modifications such as elongated appendages
While Speleosiro is monotypic Purcellia is currentlyrepresented by two additional species found in eastern SouthAfrica Purcellia transvaalica Lawrence 1963 is found inthe Hanglip forest of the Limpopo Province and has anappearance typical of other Purcellia species Purcelliaperegrinator Lawrence 1963 found in Mariepskop ForestMpumalanga Province has characteristics more typical ofthe genus Parapurcellia which has a distribution in the NatalDrakensberg and NatalndashZululand Coast of the Eastern Cape and
KwazulundashNatal Provinces In fact Lawrence (1963 279) statedthat lsquoThis species resembles [Parapurcellia] silvicola fromZululand in the shape of the tarsus IVrsquo In particular it hasozophores oriented at 45 off the lateral margin rather thandorsal ozophores projections of the anterior gonostome walla small (~25mm) body length and no evidence of functionaleyes under light microscopy ndash unlike Purcellia which haslensless eyes situated in the bases of the ozophores Based onthese characters as well as several others discussed belowand consistent placement of Purcellia peregrinator withinParapurcellia in our phylogenetic analyses we reassignPurcellia peregrinator to Parapurcellia establishing the newcombination Parapurcellia peregrinator (Lawrence 1963)leaving the number of described species in Parapurcellia atfive Parapurcellia fissa (Lawrence 1939) and Parapurcelliasilvicola (Lawrence 1939) are distinctive due to their deeplylobed posterior tergites In the remaining two speciesParapurcellia monticola (Lawrence 1939) and Parapurcelliarumpiana (Lawrence 1933) the posterior tergites are relativelyunmodified and their morphologies are typical of small pettalidsalthough the former species is distinguishable from the otherSouth African species by its tall profile
The Natal Museum of South Africa (NMSA) kindlyprovided us with undescribed cyphophthalmid specimenscollected by several arachnologists between 1939 and 1985We identified two distinct morphospecies collected in theKnysna forest that straddles the border of the Eastern andWestern Cape Provinces which appear to belong in PurcelliaAn additional four morphospecies from localities in the EasternCape and KwazulundashNatal Provinces segregated years ago byW Starega who assigned tentative names to some of the speciesappear to belong in Parapurcellia We have chosen to use hisnames as much as possible to honour his early recognition of thedifferent morphologies These are described as new speciesbelow and our placement of them into their respective generais supported by morphological phylogenetic analysis
Speleosiro argasiformisPurcellia illustrans
Parapurcellia peregrinator
Parapurcellia fissa
Purcellia griswoldi sp nov
Parapurcellia amatola sp nov
Parapurcellia staregai sp nov
Parapurcellia minuta sp novParapurcellia natalia sp nov
Purcellia lawrencei sp nov
Parapurcellia monticolaParapurcellia rumpiana
Parapurcellia silvicola
Purcellia transvaalica
NORTHERN CAPE
WESTERN CAPE
EASTERN CAPE
FREE STATE
NORTH WEST
LIMPOPO
GAU-TENG
KWAZULU-NATAL
MPUMA-LANGA
NORTHERN CAPE
WESTERN CAPE
EASTERN CAPE
FREE STATE
NORTH WEST
LIMPOPO
GAU-TENG
KWAZULU-NATAL
MPUMA-LANGA
Parapurcellia convexa sp nov
Fig 1 Mapofpettalid type collection localities inSouthAfricaCircles indicate localities of speciesdescribedbeforethiswork Stars indicate new species described herein Square indicatesParapurcellia peregrinator (Lawrence 1963)new comb whose genus is reassigned herein
372 Invertebrate Systematics B L de Bivort and G Giribet
The NMSA collections along with most collections ofpreviously described species were made before the era ofmolecular phylogenetics Their long-term storage in 70ethanol has precluded the possibility of analysing the species-level relationships of the South African pettalids using molecular
sequences In order to investigate the phylogenetic relationshipsof the SouthAfricanmaterial we chose to perform aphylogeneticanalysis under the parsimony criterion using all availablemorphological data Attempts at lsquototal evidencersquo analysis haveproved successful in past analyses of the Cyphophthalmi
Table 1 Discrete character matrixCharacter codings are explained in Appendix 2
Taxa Characters000000000 1111111111 2222222222 3333333333 4444444444 5555555555 666
123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012
Parapurcellia staregai sp nov 032200011 031101100 011001100 00-000100 0112000100 1130100100 0
Parapurcellia amatolae sp nov 032200011 031100100 1101011000 00-0020100 0112000100 1130100100 0
Parapurcellia convexa sp nov 032200011 031100100 0101011100 00-0020100 0112000110 1120100101 0
Parapurcellia natalia sp nov 032200011 031000100 011101100 00-000100 0112000110 1130100100 0
Parapurcellia minuta sp nov 032200011 031101100 0101011000 00-0020110 0112000110 10-0100100 0
Parapurcellia peregrinator 032200011 021100100 111101110 00-00300 0112000110 1120100100 0
Parapurcellia silvicola 032200011 031100000 0101011100 00-0020100 01100110 0100100101 0
Parapurcellia monticola 032200011 021100000 0101011100 00-0020100 01100110 1130100100 0
Parapurcellia rumpiana 032200011 021001100 010111101 0-30100 01100110 110100100 0
Parapurcellia fissa 032200011 031100100 0101110 30100 0100110 130100100 0
Purcellia transvaalica 222200011 020101100 110101100 00-0020 0112100100 1140110100 0
Purcellia illustrans 222200011 000100000 1101011101 00-0020100 0112100100 0140110100 001
Purcellia griswoldi sp nov 222200010 020100100 1001011100 00-0020100 0112100100 1140110100 011
Purcellia lawrencei sp nov 222200011 030100100 110101100 00-0020100 0112100100 1140110100 0
Speleosiro argasiformis 022200011 000100000 1100011101 0-0020100 0111000100 0140110100 0
Rakaia isolata 232200011 000101000 110101100 00-0020000 0100110 0100101101 0
Rakaia lindsayi 232200011 000100000 1100010210 00-0020000 01100110 0100100101 0
Rakaia magna australis 232200011 000101000 1100011200 00-0020000 01100110 0100100100 0
Rakaia media 232200011 000100000 1101011200 00-0020000 011200110 0100100100 0
Rakaia solitaria 132200011 000100000 1101011200 00-0020000 01100110 0100101100 0
Rakaia sorenseni digitata 232200011 000100000 1101011100 00-0020050 010ndashndash00110 00-01-1000 0
Aoraki crypta 232200011 000100000 1110010101 00-0020000 011200110 01311-0000 0
Aoraki denticula 232200011 000100000 1110010201 00-0021000 011200110 01301-0001 0
Aoraki inerma 232200011 000100000 1110010201 00-0020000 011200110 01311-0000 0
Aoraki longitarsa 232200011 000100000 1110010201 00-0021000 0100110 0100100101 1
Aoraki tumidata 232200011 000101000 11001011 0-2 112100111 10-0101101 0
Karripurcellia harveyi 232200011 000100010 1110010210 00-0020000 010ndashndash10100 00-00-0000 0
Karripurcellia peckorum 232200011 000101010 1110010210 00-0020000 010ndashndash10100 00-00-0000 001
Karripurcellia sierwaldae 232200011 000101010 1100011200 00-0020000 010ndashndash10100 00-00-0000 0
Austropurcellia scoparia 232200011 000100000 1101011101 00-0030100 0114000110 01101-0000 11
Austropurcellia woodwardi 232200011 020100000 111101 00-0030100 011000110 0130100100 1
Chileogovea jocasta 132200011 001100000 1110010101 00-1140000 0112100100 00-00-0000 0
Chileogovea oedipus 132100011 001100000 1110010211 00-1141000 0112100101 10-00-0000 01
Pettalus lampetides 122200011 001100010 1110110001 00-0041001 010ndashndash02100 00-01-1000 01
Pettalus cimiciformis 122100011 00310001 1110010101 00-4100 010ndashndash02100 00-01-1000 0
Troglosiro aelleni 010210011 001000000 2110000201 0110030000 001003-00 00-00-0000 0
Troglosiro longifossa 010210011 0010100100 2110000101 0110030000 001003-00 00-00-0010 0
Troglosiro ninqua 010210011 0010100000 2110000000 0110030000 001003-00 00-00-0010 0
Huitaca ventralis 010200101 111-100010 2110010201 0110030001 001003-00 00-00-0000 00
Neogovea sp 010200111 1310101010 1101000 0 003-00 00-00-0000 0
Metagovea philipi 010200111 12110000 2110000101 0110030001 001003-00 00-00-0000 001
Cyphophthalmus duricorius 010211011 0020000101 0000010100 00-0010000 001103-01 10-00-0000 011
Parasiro coiffaiti 000201010 0010000101 0100010000 0111120000 000ndashndash01000 00-00-0000 010
Siro acaroides 010211010 0010000100 0000000000 00-1110000 0011103-00 10-01-0000 011
Siro rubens 010211011 0020000100 0000020100 00-0010000 0011103-01 00-00-0000 011
Suzukielus sauteri 012200011 001000100 0000020100 1101120100 0013000101 00-00-0000 011
Stylocellus ramblae 111200101 000010001 011011021 00-0151010 020ndashndash00000 00-00-0000 0
Stylocellus tambusisi 111200100 121010001 01100102 00-001010 020ndashndash00000 10-00-0000 0
Fangensis cavernarus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001
Fangensis spelaeus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001
Systematic revision of South African pettalids Invertebrate Systematics 373
Tab
le2
Con
tinu
ouscharactermatrixas
itap
pearsin
the11
weigh
tedcombinedan
alysis
Character
definitio
nsaregivenin
App
endix3
12
34
56
78
910
1112
1314
1516
1718
1920
2122
2324
2526
2728
2930
3132
3334
3536
3738
39
Parap
urcelliastaregaisp
nov
054
059
058
052
053
053
059
047
064
051
058
063
047
067
069
075
048
071
070
056
055
049
062
073
057
057
052
052
054
066
054
068
048
048
056
056
067
061
052
Parap
urcelliaam
atolaesp
nov
047
061
057
056
056
064
052
047
065
057
053
072
051
062
061
059
062
070
066
056
042
063
062
053
053
058
058
057
072
055
050
061
043
049
056
056
066
062
049
Parapurcelliaconvexa
spn
ov
048
069
058
075
067
066
049
053
060
051
050
078
081
063
074
051
064
062
071
059
056
065
057
059
045
066
085
075
071
070
068
052
057
051
056
056
061
060
054
Parapurcellianatalia
sp
nov
044
056
057
052
053
057
049
047
057
049
060
063
048
050
071
061
105
082
074
060
053
059
061
076
059
059
052
052
065
056
043
065
043
047
056
056
071
058
049
Parapurcelliaminuta
spn
ov
042
056
056
052
053
046
050
047
057
054
081
075
047
068
072
082
048
069
062
060
059
069
059
050
052
057
052
052
043
067
055
064
057
052
056
056
067
058
048
Parapurcelliaperegrinator
059
061
058
059
055
057
059
047
077
055
065
065
048
063
049
048
066
061
057
061
055
056
057
062
071
059
060
055
058
061
053
062
053
047
056
056
065
059
059
Parapurcelliasilvicola
055
063
063
076
064
049
062
050
062
062
060
072
077
053
079
050
065
079
070
059
060
065
064
064
057
071
080
067
047
097
059
068
040
059
056
056
064
058
070
Parapurcelliamonticola
054
061
059
061
053
056
045
053
073
052
058
074
045
055
064
073
050
070
059
057
065
062
058
064
061
063
063
052
058
054
066
049
074
051
056
056
066
063
062
Parap
urcelliarumpiana
051
059
056
057
053
050
040
053
067
078
046
073
075
057
073
080
037
074
071
057
061
058
054
056
059
059
060
052
049
062
066
039
061
046
056
056
075
060
055
Parapurcelliafissa
043
064
064
052
053
056
059
053
065
052
055
070
071
069
076
063
055
066
060
057
063
079
078
060
051
085
052
052
062
036
068
083
053
053
056
056
068
060
052
Purcelliatransvaalica
069
064
057
052
053
048
060
058
061
058
041
071
070
057
074
050
061
053
072
050
067
050
056
054
056
056
052
052
046
056
054
051
050
036
056
056
074
058
048
Purcelliaillustrans
072
063
055
052
053
052
060
087
081
056
061
063
062
061
077
049
059
075
071
051
065
058
051
078
087
054
052
052
050
089
096
053
076
036
056
056
078
061
042
Purcelliagriswoldisp
nov
062
063
057
057
053
043
062
055
067
053
065
066
062
044
064
068
050
057
058
048
075
059
055
053
058
058
057
052
041
061
061
060
060
040
056
056
075
057
045
Purcellialawrenceisp
nov
062
061
056
057
053
051
066
057
057
053
059
069
067
047
054
066
053
068
067
048
080
063
052
056
050
055
057
052
050
063
068
064
068
039
056
056
075
053
044
Speleosiro
argasiform
is097
077
060
075
059
073
091
048
071
051
049
044
063
039
056
055
068
031
035
045
108
046
054
047
050
057
064
056
061
053
056
060
057
047
056
056
070
058
043
Rakaiaisolate
059
077
067
102
058
069
066
061
059
063
067
077
077
070
062
046
077
054
068
052
062
062
072
061
043
076
104
058
071
066
061
068
059
062
056
056
058
064
099
Rakaialin
dsayi
061
063
062
083
053
059
053
057
054
066
071
056
071
077
067
086
046
079
059
051
057
071
058
057
061
070
082
052
061
063
037
055
037
056
056
056
062
066
062
Rakaiamag
naau
stralis
074
068
064
078
061
085
070
047
077
085
072
052
071
057
043
053
067
068
062
054
062
062
066
068
058
060
072
059
078
065
065
059
067
053
056
056
065
058
075
Rakaiamedia
062
070
062
063
057
055
077
062
066
052
066
058
070
081
066
087
041
065
064
049
057
057
068
058
068
061
062
057
054
058
054
077
057
064
056
056
061
063
077
Rakaiasolitaria
057
065
059
071
053
065
052
060
079
053
074
080
051
055
051
046
070
065
068
053
077
043
055
037
060
065
073
052
069
069
067
055
065
066
056
056
053
062
074
Rakaiasorensenidigitata
057
071
055
054
055
075
071
080
065
066
056
093
052
085
038
054
064
069
065
051
075
075
053
066
069
054
055
056
083
060
059
080
058
074
056
056
047
053
078
Aorakicrypta
071
079
061
058
059
085
069
088
041
059
049
059
043
072
059
047
071
060
064
052
060
037
051
039
053
071
057
058
081
067
062
061
053
039
056
056
077
075
086
Aorakidenticula
057
067
055
068
062
063
048
080
054
054
056
065
051
085
058
060
067
068
071
053
066
065
051
057
069
055
071
064
066
059
063
050
063
044
075
072
071
070
070
Aorakiinerma
067
066
057
063
063
062
056
070
075
056
044
066
065
077
053
051
066
060
059
054
073
066
053
053
063
057
062
063
060
054
060
050
059
062
056
056
059
072
074
Aorakilong
itarsa
066
076
073
098
109
065
058
069
056
058
083
056
049
062
028
049
061
063
064
050
061
059
066
079
060
091
094
107
064
076
062
055
058
064
056
056
059
129
062
Aorakitumidata
064
072
058
081
069
079
070
072
063
068
071
073
079
083
054
080
060
073
070
050
062
072
055
054
062
060
083
070
081
085
062
070
050
070
075
096
080
059
071
Karripurcellia
harveyi
067
052
058
061
066
051
079
089
062
052
069
056
073
060
074
075
037
050
043
055
056
063
060
069
091
052
060
065
049
066
050
072
048
067
056
056
047
057
058
Karripurcellia
peckorum
068
062
057
071
068
069
062
070
051
094
057
064
070
057
052
045
071
046
037
055
060
064
057
059
076
054
069
067
064
070
046
053
037
068
056
056
047
056
056
Karripurcellia
sierwaldae
056
056
057
052
053
042
062
061
067
088
056
061
076
063
070
057
060
043
032
055
062
069
062
067
068
053
052
052
039
067
057
064
051
073
064
087
047
057
060
Austropurcelliascoparia
055
050
054
052
053
067
052
068
054
051
045
065
050
064
057
044
071
065
074
059
071
072
051
054
084
053
052
052
071
052
082
055
088
062
056
056
061
091
053
Austrop
urcelliawoo
dwardi
054
065
058
052
058
056
061
069
053
053
042
066
049
066
068
053
057
062
068
054
063
047
049
036
050
067
052
060
056
065
079
064
068
059
056
056
047
061
053
Chileogovea
jocasta
049
053
055
052
054
058
050
057
055
049
047
049
058
060
055
063
050
050
058
051
074
055
053
053
060
053
052
055
061
057
069
057
068
066
082
088
082
053
069
Chileog
ovea
Oedipus
070
053
054
052
053
064
070
074
057
062
053
052
043
064
054
051
058
072
078
053
071
012
049
043
054
052
052
052
061
052
063
061
070
062
092
091
083
051
048
Pettaluslampetid
es063
066
078
052
092
067
057
058
053
055
063
039
048
055
057
063
064
068
068
046
064
050
095
090
044
084
052
093
065
042
050
052
066
074
056
056
047
052
055
Pettaluscimicifo
rmis
100
110
140
066
095
089
082
076
066
058
074
033
039
021
057
058
058
056
046
041
056
085
125
064
044
116
059
074
070
039
047
055
071
068
056
056
047
050
061
Troglosiroaelleni
059
053
059
054
066
066
064
057
054
053
052
040
067
066
072
060
064
044
053
076
045
064
066
075
062
054
054
067
068
054
061
066
064
074
056
056
047
055
058
Troglosirolong
ifossa
051
045
055
052
073
064
048
070
049
060
057
042
057
053
069
043
069
053
064
079
041
059
055
061
066
052
052
081
070
049
049
052
055
074
056
056
047
056
053
Troglosironinq
ua059
045
057
053
086
068
049
074
049
066
048
044
074
054
064
052
063
048
059
076
044
054
063
072
068
052
054
092
069
062
076
048
070
074
056
056
047
057
072
Huitaca
ventralis
075
045
057
052
060
036
084
047
105
094
063
054
058
067
072
068
054
040
056
075
057
046
058
070
053
052
052
058
039
045
038
057
047
074
056
056
047
052
048
Neogo
veasp
067
056
058
065
070
031
070
075
081
099
042
055
067
047
063
068
042
031
045
074
050
038
061
073
061
052
063
069
033
035
035
055
044
074
056
056
047
050
045
Metagovea
philipi
045
045
054
052
053
041
045
057
065
055
056
056
066
051
053
054
056
048
062
078
057
057
053
050
031
052
052
052
036
042
056
048
073
074
056
056
047
053
068
Cypho
phthalmus
duricorius
050
045
055
052
053
058
041
050
039
060
067
044
044
042
027
050
072
058
065
078
039
065
057
071
065
052
052
052
062
059
067
044
072
074
056
056
047
056
065
Parasirocoiffaiti
044
045
055
054
055
058
056
047
050
052
079
050
041
060
056
080
051
064
061
086
042
064
060
062
058
052
056
057
068
053
052
081
063
068
056
056
055
053
064
Siro
acaroids
046
055
054
052
053
055
053
049
044
055
047
051
067
060
070
051
057
058
071
069
048
068
053
044
061
053
052
052
058
062
061
062
055
051
056
056
071
055
068
Siro
rubens
049
045
056
052
053
042
077
050
041
055
065
039
057
053
062
082
037
047
052
072
035
073
059
052
086
052
052
052
036
054
085
102
100
066
119
102
056
050
049
Suzukielus
sauteri
059
045
056
052
053
060
062
048
050
051
063
068
069
065
064
066
056
082
071
075
048
071
056
067
057
052
052
052
062
079
072
066
063
071
098
091
047
056
061
Stylocellusramblae
052
045
055
052
053
064
050
060
052
048
083
058
047
056
053
055
076
045
046
092
068
060
054
046
048
052
052
052
067
053
058
053
074
074
056
056
047
053
060
Stylocellustambusisi
074
045
057
052
053
066
049
059
046
067
058
052
050
041
029
059
076
047
056
082
066
060
057
070
056
052
052
052
061
047
055
041
065
074
056
056
047
052
062
Fan
gensiscavernarus
081
064
065
052
053
073
046
047
049
065
080
054
076
053
065
048
079
048
026
075
061
073
065
078
081
059
052
052
066
063
049
038
051
074
056
056
047
053
048
Fan
gensisspelaeus
073
045
057
052
053
078
093
047
050
061
093
065
077
056
064
076
053
068
030
076
054
069
056
069
066
052
052
052
076
068
055
086
061
074
056
056
047
057
054
374 Invertebrate Systematics B L de Bivort and G Giribet
4041
4243
4445
4647
4849
5051
5253
5455
5657
5859
6061
6263
6465
6667
6869
7071
7273
7475
7677
78
Parap
urcelliastaregaisp
nov
054
049
048
053
061
054
046
048
065
057
060
056
057
059
069
055
074
063
048
049
055
058
069
062
068
080
057
066
053
057
056
052
068
059
066
054
070
051
062
Parap
urcelliaam
atolaesp
nov
047
058
055
044
066
054
046
050
081
058
068
069
065
064
068
055
068
055
049
032
047
064
064
070
066
059
053
073
046
043
035
075
066
053
068
048
065
062
056
Parapurcelliaconvexa
spn
ov
036
030
068
095
053
054
055
050
064
057
053
052
056
065
070
055
069
066
055
062
053
069
052
075
070
059
063
067
049
055
069
062
077
034
062
057
071
052
062
Parapurcellianatalia
sp
nov
050
046
050
051
054
054
046
046
079
060
056
066
072
063
064
055
062
060
057
058
046
068
053
063
063
069
042
063
045
054
073
050
062
073
056
054
060
048
054
Parap
urcelliaminuta
spn
ov
046
046
054
041
030
054
047
049
075
060
059
050
058
059
066
055
062
062
057
040
062
066
072
061
062
076
044
064
052
056
058
076
095
069
053
058
062
057
054
Parap
urcelliaperegrinator
058
058
058
050
049
054
046
054
071
063
049
056
070
062
060
055
062
060
057
070
053
063
073
077
079
056
052
078
049
061
058
057
064
079
052
048
066
055
055
Parapurcelliasilvicola
052
054
062
050
044
054
050
048
064
058
063
059
055
062
069
055
064
061
052
051
062
067
072
070
066
019
045
064
059
059
048
061
074
066
065
073
070
064
057
Parapurcelliamonticola
067
073
086
060
055
054
053
051
066
057
050
062
071
063
070
055
060
065
058
053
044
064
061
039
072
075
047
080
048
061
056
047
066
061
066
047
061
053
056
Parap
urcelliarumpian
a040
035
072
081
058
054
055
045
075
057
048
067
074
063
065
055
082
059
056
072
052
066
051
064
073
068
049
076
073
066
056
058
070
035
054
090
078
093
058
Parapurcelliafissa
060
057
065
048
059
054
057
060
063
057
049
056
066
063
060
055
062
061
051
064
056
065
062
038
074
051
057
079
050
059
043
048
069
061
067
051
071
055
060
Purcelliatransvaalica
064
068
060
051
053
054
056
057
062
055
044
054
065
055
069
055
077
068
052
054
044
068
069
066
064
058
060
058
060
071
050
043
054
051
073
055
054
056
058
Purcelliaillustrans
111
080
101
074
037
054
082
067
066
052
047
046
060
066
057
055
056
056
055
047
052
066
063
057
066
048
055
077
052
068
051
043
059
048
053
047
072
054
059
Purcelliagriswoldisp
nov
067
078
063
048
049
054
054
049
065
057
045
050
066
061
065
055
053
061
059
046
050
063
074
056
070
071
070
066
058
070
035
050
051
069
049
058
077
052
056
Purcellialawrenceisp
nov
071
081
067
052
046
054
056
052
089
054
046
040
054
061
066
055
059
059
069
047
038
074
067
054
072
068
056
056
056
065
051
050
050
050
042
062
068
052
057
Speleosiro
argasiform
is057
056
054
045
060
054
047
044
055
066
041
058
078
061
045
055
058
061
057
044
076
049
090
079
055
054
085
058
077
089
050
052
051
070
043
041
050
048
060
Rakaiaisolate
066
075
061
060
062
054
061
079
032
073
059
069
071
063
059
055
066
060
066
044
061
052
039
053
069
056
038
058
049
050
049
077
071
058
077
062
065
064
059
Rakaialin
dsayi
050
060
058
056
068
054
057
054
070
057
067
065
060
067
060
055
055
056
068
059
073
059
060
044
078
074
049
053
060
059
050
064
057
085
059
055
064
066
058
Rakaiamag
naau
stralis
060
047
055
055
055
054
046
052
059
078
064
074
072
066
054
055
061
064
073
060
060
064
050
056
070
047
058
068
062
052
053
074
049
049
058
061
070
077
058
Rakaiamedia
060
082
041
052
062
054
061
052
065
078
064
082
075
061
058
055
060
062
075
060
047
059
043
059
063
071
060
064
049
037
076
056
070
050
061
063
070
051
060
Rakaiasolitaria
065
071
056
052
051
054
060
066
068
067
074
075
063
069
053
055
078
056
065
060
053
064
047
055
063
051
057
081
050
048
074
072
056
065
075
069
073
052
058
Rakaiasorensenidigitata
076
071
066
060
034
054
087
052
064
060
049
030
039
084
055
055
065
065
065
060
068
057
072
072
072
051
067
069
061
062
061
078
076
078
077
073
071
070
048
Aorakicrypta
071
069
053
060
054
054
082
048
068
067
068
063
051
069
055
055
059
073
071
054
064
047
075
078
071
056
080
040
068
049
067
081
042
064
068
041
047
057
048
Aorakidenticula
075
068
054
061
068
054
085
055
052
057
068
055
048
077
054
055
070
060
056
065
047
073
071
049
074
066
053
055
057
035
085
061
070
067
058
040
059
054
060
Aorakiinerma
069
070
071
078
065
054
069
046
068
060
064
063
052
072
050
055
081
064
054
050
058
060
062
072
042
055
057
073
070
068
054
052
047
043
079
046
057
055
061
Aorakilong
itarsa
069
066
057
062
073
054
067
078
064
059
075
063
056
071
056
055
065
073
057
069
073
043
074
071
064
050
071
054
064
049
071
078
050
052
062
056
063
057
062
Aorakitumidata
066
062
067
047
063
054
073
062
066
055
074
087
071
070
065
055
051
065
072
062
095
023
074
082
079
081
077
063
082
060
064
105
068
085
055
059
071
069
069
Karripurcellia
harveyi
050
061
072
074
086
054
084
076
057
054
047
047
064
062
057
055
056
054
061
077
075
053
060
073
053
047
066
059
064
070
054
051
055
061
045
052
054
050
060
Karripurcellia
peckorum
046
050
066
077
087
054
067
056
050
067
045
047
062
064
056
055
059
059
074
071
069
056
061
059
063
045
055
049
082
067
054
055
042
054
081
071
051
090
055
Karripurcellia
sierwaldae
050
052
064
067
068
054
062
061
060
050
060
044
044
064
065
055
052
060
051
060
060
058
057
064
061
060
063
070
079
074
051
040
045
058
056
081
057
097
048
Austropurcelliascoparia
079
078
069
066
053
054
074
064
051
052
080
056
037
089
038
055
056
057
080
050
057
076
067
062
060
048
053
056
052
062
064
060
061
068
062
066
062
052
063
Austrop
urcelliawoo
dwardi
072
067
074
061
054
054
075
079
049
062
075
067
047
074
036
055
058
058
053
054
051
086
061
052
056
059
038
054
054
065
060
046
058
049
063
080
067
054
048
Chileogovea
jocasta
067
064
065
054
054
054
061
052
054
055
069
066
052
069
045
055
049
058
057
057
063
066
042
053
059
068
068
059
056
074
054
056
057
060
043
057
046
050
048
Chileog
ovea
oedipu
s064
068
070
070
048
054
070
076
081
056
070
056
046
069
056
055
054
059
054
085
072
060
047
044
076
057
066
055
072
074
048
064
054
076
046
052
041
060
048
Pettaluslampetid
es063
055
059
058
058
054
057
051
063
063
053
077
084
052
075
055
058
051
057
058
054
052
051
057
060
063
062
052
073
067
054
051
039
075
083
040
034
053
061
Pettaluscimicifo
rmis
063
061
050
062
048
062
060
058
044
065
044
054
079
048
070
055
042
048
069
049
060
051
057
051
058
062
082
054
051
061
077
059
060
053
072
046
062
051
057
Troglosiroaelleni
059
056
029
089
066
085
058
054
066
046
078
071
050
062
045
090
056
060
068
076
055
055
066
058
045
064
076
054
064
065
059
049
055
055
046
049
037
052
060
Troglosirolongifo
ssa
059
055
058
070
077
085
077
053
061
045
069
071
058
048
054
093
038
065
070
085
059
056
052
082
044
051
071
049
056
045
075
053
050
053
038
063
073
064
048
Troglosironinq
ua066
051
072
073
079
083
077
061
060
041
068
067
053
045
064
092
062
059
058
067
059
053
070
075
052
061
082
049
075
076
046
055
048
051
036
053
045
068
048
Huitaca
ventralis
045
045
040
050
059
085
046
058
056
041
049
070
083
032
075
081
031
044
049
051
062
061
070
030
046
062
071
090
108
096
085
062
043
073
043
063
033
084
048
Neogo
veasp
042
067
051
086
073
087
071
066
031
046
039
066
087
037
059
055
066
130
104
061
066
071
062
039
042
072
075
076
074
077
079
055
050
036
084
064
038
095
061
Metag
ovea
philipi
057
068
061
073
063
085
062
044
040
041
066
069
059
039
057
055
061
044
080
064
056
085
058
045
035
059
041
075
059
063
053
049
046
047
065
054
034
059
048
Cypho
phthalmus
duricorius
060
046
053
056
061
086
050
058
065
056
053
064
073
054
051
087
060
064
057
082
060
061
060
065
048
047
064
036
055
052
070
059
066
065
071
077
072
064
078
Parasirocoiffaiti
056
064
071
050
066
081
046
074
063
060
089
070
051
051
055
078
055
065
053
062
063
060
032
073
042
079
053
052
044
048
078
068
088
061
060
077
070
055
048
Siro
acaroides
052
040
038
053
081
085
049
077
058
060
057
079
073
055
059
096
032
061
049
068
045
071
040
057
055
059
058
043
065
064
063
065
066
050
059
067
056
059
048
Siro
rubens
063
050
064
053
063
085
050
069
056
049
076
069
054
060
051
087
073
067
078
089
056
064
034
060
046
084
064
038
056
054
068
064
076
049
045
063
068
058
073
Suzukielus
sauteri
064
047
063
048
073
054
047
101
069
053
063
050
048
052
075
055
060
062
067
077
059
058
042
069
068
069
052
050
049
048
075
061
066
056
069
065
074
050
068
Stylocellusramblae
060
072
062
053
058
054
062
073
041
098
074
042
040
035
071
055
025
044
038
053
105
053
072
051
030
061
082
052
056
052
067
075
059
072
063
066
048
047
099
Stylocellustambusisi
056
063
061
052
060
054
056
069
047
096
073
050
041
040
067
055
063
044
038
042
064
060
074
050
036
048
063
044
060
049
067
060
058
078
057
085
046
062
087
Fan
gensiscavernarus
047
044
040
056
075
054
046
070
036
085
050
042
058
041
084
055
077
044
038
066
076
035
052
064
049
055
070
046
051
044
056
067
062
071
050
058
060
057
096
Fan
gensisspelaeus
049
061
044
055
079
054
046
070
047
086
046
035
058
042
105
055
087
044
038
065
077
029
068
065
060
084
050
047
051
045
078
070
083
057
054
071
065
056
096
Systematic revision of South African pettalids Invertebrate Systematics 375
combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)
Methods and abbreviations
Specimens used in this study were provided by the followinginstitutions
AMNH American Museum of Natural History New York(USA)
BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard
University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand
Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo
Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg
(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg
Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)
Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg
(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen
Copenhagen (Denmark)
Specimen handling and illustration
In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was
photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy
Character coding
Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3
Phylogenetic analyses
The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)
The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the
376 Invertebrate Systematics B L de Bivort and G Giribet
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
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trict
consensusof
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underparsim
onyanalysisof
thecontinuous
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pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
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phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
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81
41
21
11
12
14
18
Pet
talid
ae S
tylo
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Neo
gove
idae
Tro
glos
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(T
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Neo
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)K
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trop
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ogov
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(Pur
celli
a gr
isw
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renc
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p n
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(Pet
talid
ae +
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ukie
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(Aor
aki +
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(Pet
talid
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nida
e)
(Kar
ripur
celli
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talu
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(Par
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celli
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leos
iro)
(Pur
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Aus
trop
urce
llia)
AB
C
Ao
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cula
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Tro
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a Ka
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arv
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99 51 69
75 92 95 8581
80 69
94
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74 83
86 86 5665
6379
8877 100
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81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
2007 2009 de Bivort et al 2010) Molecular evidence favoursthe monophyly of all genera as currently defined although only afew (Aoraki Rakaia Pettalus) have been thoroughly sampled inmolecular studies (Boyer and Giribet 2007 Boyer et al 2007a2007b) In these studies the African pettalids were restricted torepresentatives of only three species one in the genus Purcelliaand two in the genus Parapurcellia and South Africa wasparaphyletic with respect to other landmasses
The described South African pettalid fauna is currentlydistributed in two geographic regions (Fig 1) Two speciesare found in the Western Cape province specifically in andaround Table Mountain Purcellia illustrans Hansen ampSoslashrensen 1904 has been collected in forests surroundingTable Mountain Speleosiro argasiformis Lawrence 1931 isa large (5mm) cyphophthalmid found in the Wynberg cavesystem in Table Mountain (Juberthie 1970) The formerspecies is a medium-sized cyphophthalmid with morphologicalcharacteristics typical of other members of South Africanpettalids such as relatively unmodified opisthosomal posteriortergites in males and females second palp articles that are lessthan half ornamented and bilobed anal plates in males Forillustrations of these and other cyphophthalmid morphologicalterms see de Bivort et al (2010) One conspicuous featureof P illustrans is the position of its anal plate which is shiftedanteriorly onto the ventral surface of the animal comparedwith other pettalids Speleosiro argasiformis has conspicuoustroglomorphic modifications such as elongated appendages
While Speleosiro is monotypic Purcellia is currentlyrepresented by two additional species found in eastern SouthAfrica Purcellia transvaalica Lawrence 1963 is found inthe Hanglip forest of the Limpopo Province and has anappearance typical of other Purcellia species Purcelliaperegrinator Lawrence 1963 found in Mariepskop ForestMpumalanga Province has characteristics more typical ofthe genus Parapurcellia which has a distribution in the NatalDrakensberg and NatalndashZululand Coast of the Eastern Cape and
KwazulundashNatal Provinces In fact Lawrence (1963 279) statedthat lsquoThis species resembles [Parapurcellia] silvicola fromZululand in the shape of the tarsus IVrsquo In particular it hasozophores oriented at 45 off the lateral margin rather thandorsal ozophores projections of the anterior gonostome walla small (~25mm) body length and no evidence of functionaleyes under light microscopy ndash unlike Purcellia which haslensless eyes situated in the bases of the ozophores Based onthese characters as well as several others discussed belowand consistent placement of Purcellia peregrinator withinParapurcellia in our phylogenetic analyses we reassignPurcellia peregrinator to Parapurcellia establishing the newcombination Parapurcellia peregrinator (Lawrence 1963)leaving the number of described species in Parapurcellia atfive Parapurcellia fissa (Lawrence 1939) and Parapurcelliasilvicola (Lawrence 1939) are distinctive due to their deeplylobed posterior tergites In the remaining two speciesParapurcellia monticola (Lawrence 1939) and Parapurcelliarumpiana (Lawrence 1933) the posterior tergites are relativelyunmodified and their morphologies are typical of small pettalidsalthough the former species is distinguishable from the otherSouth African species by its tall profile
The Natal Museum of South Africa (NMSA) kindlyprovided us with undescribed cyphophthalmid specimenscollected by several arachnologists between 1939 and 1985We identified two distinct morphospecies collected in theKnysna forest that straddles the border of the Eastern andWestern Cape Provinces which appear to belong in PurcelliaAn additional four morphospecies from localities in the EasternCape and KwazulundashNatal Provinces segregated years ago byW Starega who assigned tentative names to some of the speciesappear to belong in Parapurcellia We have chosen to use hisnames as much as possible to honour his early recognition of thedifferent morphologies These are described as new speciesbelow and our placement of them into their respective generais supported by morphological phylogenetic analysis
Speleosiro argasiformisPurcellia illustrans
Parapurcellia peregrinator
Parapurcellia fissa
Purcellia griswoldi sp nov
Parapurcellia amatola sp nov
Parapurcellia staregai sp nov
Parapurcellia minuta sp novParapurcellia natalia sp nov
Purcellia lawrencei sp nov
Parapurcellia monticolaParapurcellia rumpiana
Parapurcellia silvicola
Purcellia transvaalica
NORTHERN CAPE
WESTERN CAPE
EASTERN CAPE
FREE STATE
NORTH WEST
LIMPOPO
GAU-TENG
KWAZULU-NATAL
MPUMA-LANGA
NORTHERN CAPE
WESTERN CAPE
EASTERN CAPE
FREE STATE
NORTH WEST
LIMPOPO
GAU-TENG
KWAZULU-NATAL
MPUMA-LANGA
Parapurcellia convexa sp nov
Fig 1 Mapofpettalid type collection localities inSouthAfricaCircles indicate localities of speciesdescribedbeforethiswork Stars indicate new species described herein Square indicatesParapurcellia peregrinator (Lawrence 1963)new comb whose genus is reassigned herein
372 Invertebrate Systematics B L de Bivort and G Giribet
The NMSA collections along with most collections ofpreviously described species were made before the era ofmolecular phylogenetics Their long-term storage in 70ethanol has precluded the possibility of analysing the species-level relationships of the South African pettalids using molecular
sequences In order to investigate the phylogenetic relationshipsof the SouthAfricanmaterial we chose to perform aphylogeneticanalysis under the parsimony criterion using all availablemorphological data Attempts at lsquototal evidencersquo analysis haveproved successful in past analyses of the Cyphophthalmi
Table 1 Discrete character matrixCharacter codings are explained in Appendix 2
Taxa Characters000000000 1111111111 2222222222 3333333333 4444444444 5555555555 666
123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012
Parapurcellia staregai sp nov 032200011 031101100 011001100 00-000100 0112000100 1130100100 0
Parapurcellia amatolae sp nov 032200011 031100100 1101011000 00-0020100 0112000100 1130100100 0
Parapurcellia convexa sp nov 032200011 031100100 0101011100 00-0020100 0112000110 1120100101 0
Parapurcellia natalia sp nov 032200011 031000100 011101100 00-000100 0112000110 1130100100 0
Parapurcellia minuta sp nov 032200011 031101100 0101011000 00-0020110 0112000110 10-0100100 0
Parapurcellia peregrinator 032200011 021100100 111101110 00-00300 0112000110 1120100100 0
Parapurcellia silvicola 032200011 031100000 0101011100 00-0020100 01100110 0100100101 0
Parapurcellia monticola 032200011 021100000 0101011100 00-0020100 01100110 1130100100 0
Parapurcellia rumpiana 032200011 021001100 010111101 0-30100 01100110 110100100 0
Parapurcellia fissa 032200011 031100100 0101110 30100 0100110 130100100 0
Purcellia transvaalica 222200011 020101100 110101100 00-0020 0112100100 1140110100 0
Purcellia illustrans 222200011 000100000 1101011101 00-0020100 0112100100 0140110100 001
Purcellia griswoldi sp nov 222200010 020100100 1001011100 00-0020100 0112100100 1140110100 011
Purcellia lawrencei sp nov 222200011 030100100 110101100 00-0020100 0112100100 1140110100 0
Speleosiro argasiformis 022200011 000100000 1100011101 0-0020100 0111000100 0140110100 0
Rakaia isolata 232200011 000101000 110101100 00-0020000 0100110 0100101101 0
Rakaia lindsayi 232200011 000100000 1100010210 00-0020000 01100110 0100100101 0
Rakaia magna australis 232200011 000101000 1100011200 00-0020000 01100110 0100100100 0
Rakaia media 232200011 000100000 1101011200 00-0020000 011200110 0100100100 0
Rakaia solitaria 132200011 000100000 1101011200 00-0020000 01100110 0100101100 0
Rakaia sorenseni digitata 232200011 000100000 1101011100 00-0020050 010ndashndash00110 00-01-1000 0
Aoraki crypta 232200011 000100000 1110010101 00-0020000 011200110 01311-0000 0
Aoraki denticula 232200011 000100000 1110010201 00-0021000 011200110 01301-0001 0
Aoraki inerma 232200011 000100000 1110010201 00-0020000 011200110 01311-0000 0
Aoraki longitarsa 232200011 000100000 1110010201 00-0021000 0100110 0100100101 1
Aoraki tumidata 232200011 000101000 11001011 0-2 112100111 10-0101101 0
Karripurcellia harveyi 232200011 000100010 1110010210 00-0020000 010ndashndash10100 00-00-0000 0
Karripurcellia peckorum 232200011 000101010 1110010210 00-0020000 010ndashndash10100 00-00-0000 001
Karripurcellia sierwaldae 232200011 000101010 1100011200 00-0020000 010ndashndash10100 00-00-0000 0
Austropurcellia scoparia 232200011 000100000 1101011101 00-0030100 0114000110 01101-0000 11
Austropurcellia woodwardi 232200011 020100000 111101 00-0030100 011000110 0130100100 1
Chileogovea jocasta 132200011 001100000 1110010101 00-1140000 0112100100 00-00-0000 0
Chileogovea oedipus 132100011 001100000 1110010211 00-1141000 0112100101 10-00-0000 01
Pettalus lampetides 122200011 001100010 1110110001 00-0041001 010ndashndash02100 00-01-1000 01
Pettalus cimiciformis 122100011 00310001 1110010101 00-4100 010ndashndash02100 00-01-1000 0
Troglosiro aelleni 010210011 001000000 2110000201 0110030000 001003-00 00-00-0000 0
Troglosiro longifossa 010210011 0010100100 2110000101 0110030000 001003-00 00-00-0010 0
Troglosiro ninqua 010210011 0010100000 2110000000 0110030000 001003-00 00-00-0010 0
Huitaca ventralis 010200101 111-100010 2110010201 0110030001 001003-00 00-00-0000 00
Neogovea sp 010200111 1310101010 1101000 0 003-00 00-00-0000 0
Metagovea philipi 010200111 12110000 2110000101 0110030001 001003-00 00-00-0000 001
Cyphophthalmus duricorius 010211011 0020000101 0000010100 00-0010000 001103-01 10-00-0000 011
Parasiro coiffaiti 000201010 0010000101 0100010000 0111120000 000ndashndash01000 00-00-0000 010
Siro acaroides 010211010 0010000100 0000000000 00-1110000 0011103-00 10-01-0000 011
Siro rubens 010211011 0020000100 0000020100 00-0010000 0011103-01 00-00-0000 011
Suzukielus sauteri 012200011 001000100 0000020100 1101120100 0013000101 00-00-0000 011
Stylocellus ramblae 111200101 000010001 011011021 00-0151010 020ndashndash00000 00-00-0000 0
Stylocellus tambusisi 111200100 121010001 01100102 00-001010 020ndashndash00000 10-00-0000 0
Fangensis cavernarus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001
Fangensis spelaeus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001
Systematic revision of South African pettalids Invertebrate Systematics 373
Tab
le2
Con
tinu
ouscharactermatrixas
itap
pearsin
the11
weigh
tedcombinedan
alysis
Character
definitio
nsaregivenin
App
endix3
12
34
56
78
910
1112
1314
1516
1718
1920
2122
2324
2526
2728
2930
3132
3334
3536
3738
39
Parap
urcelliastaregaisp
nov
054
059
058
052
053
053
059
047
064
051
058
063
047
067
069
075
048
071
070
056
055
049
062
073
057
057
052
052
054
066
054
068
048
048
056
056
067
061
052
Parap
urcelliaam
atolaesp
nov
047
061
057
056
056
064
052
047
065
057
053
072
051
062
061
059
062
070
066
056
042
063
062
053
053
058
058
057
072
055
050
061
043
049
056
056
066
062
049
Parapurcelliaconvexa
spn
ov
048
069
058
075
067
066
049
053
060
051
050
078
081
063
074
051
064
062
071
059
056
065
057
059
045
066
085
075
071
070
068
052
057
051
056
056
061
060
054
Parapurcellianatalia
sp
nov
044
056
057
052
053
057
049
047
057
049
060
063
048
050
071
061
105
082
074
060
053
059
061
076
059
059
052
052
065
056
043
065
043
047
056
056
071
058
049
Parapurcelliaminuta
spn
ov
042
056
056
052
053
046
050
047
057
054
081
075
047
068
072
082
048
069
062
060
059
069
059
050
052
057
052
052
043
067
055
064
057
052
056
056
067
058
048
Parapurcelliaperegrinator
059
061
058
059
055
057
059
047
077
055
065
065
048
063
049
048
066
061
057
061
055
056
057
062
071
059
060
055
058
061
053
062
053
047
056
056
065
059
059
Parapurcelliasilvicola
055
063
063
076
064
049
062
050
062
062
060
072
077
053
079
050
065
079
070
059
060
065
064
064
057
071
080
067
047
097
059
068
040
059
056
056
064
058
070
Parapurcelliamonticola
054
061
059
061
053
056
045
053
073
052
058
074
045
055
064
073
050
070
059
057
065
062
058
064
061
063
063
052
058
054
066
049
074
051
056
056
066
063
062
Parap
urcelliarumpiana
051
059
056
057
053
050
040
053
067
078
046
073
075
057
073
080
037
074
071
057
061
058
054
056
059
059
060
052
049
062
066
039
061
046
056
056
075
060
055
Parapurcelliafissa
043
064
064
052
053
056
059
053
065
052
055
070
071
069
076
063
055
066
060
057
063
079
078
060
051
085
052
052
062
036
068
083
053
053
056
056
068
060
052
Purcelliatransvaalica
069
064
057
052
053
048
060
058
061
058
041
071
070
057
074
050
061
053
072
050
067
050
056
054
056
056
052
052
046
056
054
051
050
036
056
056
074
058
048
Purcelliaillustrans
072
063
055
052
053
052
060
087
081
056
061
063
062
061
077
049
059
075
071
051
065
058
051
078
087
054
052
052
050
089
096
053
076
036
056
056
078
061
042
Purcelliagriswoldisp
nov
062
063
057
057
053
043
062
055
067
053
065
066
062
044
064
068
050
057
058
048
075
059
055
053
058
058
057
052
041
061
061
060
060
040
056
056
075
057
045
Purcellialawrenceisp
nov
062
061
056
057
053
051
066
057
057
053
059
069
067
047
054
066
053
068
067
048
080
063
052
056
050
055
057
052
050
063
068
064
068
039
056
056
075
053
044
Speleosiro
argasiform
is097
077
060
075
059
073
091
048
071
051
049
044
063
039
056
055
068
031
035
045
108
046
054
047
050
057
064
056
061
053
056
060
057
047
056
056
070
058
043
Rakaiaisolate
059
077
067
102
058
069
066
061
059
063
067
077
077
070
062
046
077
054
068
052
062
062
072
061
043
076
104
058
071
066
061
068
059
062
056
056
058
064
099
Rakaialin
dsayi
061
063
062
083
053
059
053
057
054
066
071
056
071
077
067
086
046
079
059
051
057
071
058
057
061
070
082
052
061
063
037
055
037
056
056
056
062
066
062
Rakaiamag
naau
stralis
074
068
064
078
061
085
070
047
077
085
072
052
071
057
043
053
067
068
062
054
062
062
066
068
058
060
072
059
078
065
065
059
067
053
056
056
065
058
075
Rakaiamedia
062
070
062
063
057
055
077
062
066
052
066
058
070
081
066
087
041
065
064
049
057
057
068
058
068
061
062
057
054
058
054
077
057
064
056
056
061
063
077
Rakaiasolitaria
057
065
059
071
053
065
052
060
079
053
074
080
051
055
051
046
070
065
068
053
077
043
055
037
060
065
073
052
069
069
067
055
065
066
056
056
053
062
074
Rakaiasorensenidigitata
057
071
055
054
055
075
071
080
065
066
056
093
052
085
038
054
064
069
065
051
075
075
053
066
069
054
055
056
083
060
059
080
058
074
056
056
047
053
078
Aorakicrypta
071
079
061
058
059
085
069
088
041
059
049
059
043
072
059
047
071
060
064
052
060
037
051
039
053
071
057
058
081
067
062
061
053
039
056
056
077
075
086
Aorakidenticula
057
067
055
068
062
063
048
080
054
054
056
065
051
085
058
060
067
068
071
053
066
065
051
057
069
055
071
064
066
059
063
050
063
044
075
072
071
070
070
Aorakiinerma
067
066
057
063
063
062
056
070
075
056
044
066
065
077
053
051
066
060
059
054
073
066
053
053
063
057
062
063
060
054
060
050
059
062
056
056
059
072
074
Aorakilong
itarsa
066
076
073
098
109
065
058
069
056
058
083
056
049
062
028
049
061
063
064
050
061
059
066
079
060
091
094
107
064
076
062
055
058
064
056
056
059
129
062
Aorakitumidata
064
072
058
081
069
079
070
072
063
068
071
073
079
083
054
080
060
073
070
050
062
072
055
054
062
060
083
070
081
085
062
070
050
070
075
096
080
059
071
Karripurcellia
harveyi
067
052
058
061
066
051
079
089
062
052
069
056
073
060
074
075
037
050
043
055
056
063
060
069
091
052
060
065
049
066
050
072
048
067
056
056
047
057
058
Karripurcellia
peckorum
068
062
057
071
068
069
062
070
051
094
057
064
070
057
052
045
071
046
037
055
060
064
057
059
076
054
069
067
064
070
046
053
037
068
056
056
047
056
056
Karripurcellia
sierwaldae
056
056
057
052
053
042
062
061
067
088
056
061
076
063
070
057
060
043
032
055
062
069
062
067
068
053
052
052
039
067
057
064
051
073
064
087
047
057
060
Austropurcelliascoparia
055
050
054
052
053
067
052
068
054
051
045
065
050
064
057
044
071
065
074
059
071
072
051
054
084
053
052
052
071
052
082
055
088
062
056
056
061
091
053
Austrop
urcelliawoo
dwardi
054
065
058
052
058
056
061
069
053
053
042
066
049
066
068
053
057
062
068
054
063
047
049
036
050
067
052
060
056
065
079
064
068
059
056
056
047
061
053
Chileogovea
jocasta
049
053
055
052
054
058
050
057
055
049
047
049
058
060
055
063
050
050
058
051
074
055
053
053
060
053
052
055
061
057
069
057
068
066
082
088
082
053
069
Chileog
ovea
Oedipus
070
053
054
052
053
064
070
074
057
062
053
052
043
064
054
051
058
072
078
053
071
012
049
043
054
052
052
052
061
052
063
061
070
062
092
091
083
051
048
Pettaluslampetid
es063
066
078
052
092
067
057
058
053
055
063
039
048
055
057
063
064
068
068
046
064
050
095
090
044
084
052
093
065
042
050
052
066
074
056
056
047
052
055
Pettaluscimicifo
rmis
100
110
140
066
095
089
082
076
066
058
074
033
039
021
057
058
058
056
046
041
056
085
125
064
044
116
059
074
070
039
047
055
071
068
056
056
047
050
061
Troglosiroaelleni
059
053
059
054
066
066
064
057
054
053
052
040
067
066
072
060
064
044
053
076
045
064
066
075
062
054
054
067
068
054
061
066
064
074
056
056
047
055
058
Troglosirolong
ifossa
051
045
055
052
073
064
048
070
049
060
057
042
057
053
069
043
069
053
064
079
041
059
055
061
066
052
052
081
070
049
049
052
055
074
056
056
047
056
053
Troglosironinq
ua059
045
057
053
086
068
049
074
049
066
048
044
074
054
064
052
063
048
059
076
044
054
063
072
068
052
054
092
069
062
076
048
070
074
056
056
047
057
072
Huitaca
ventralis
075
045
057
052
060
036
084
047
105
094
063
054
058
067
072
068
054
040
056
075
057
046
058
070
053
052
052
058
039
045
038
057
047
074
056
056
047
052
048
Neogo
veasp
067
056
058
065
070
031
070
075
081
099
042
055
067
047
063
068
042
031
045
074
050
038
061
073
061
052
063
069
033
035
035
055
044
074
056
056
047
050
045
Metagovea
philipi
045
045
054
052
053
041
045
057
065
055
056
056
066
051
053
054
056
048
062
078
057
057
053
050
031
052
052
052
036
042
056
048
073
074
056
056
047
053
068
Cypho
phthalmus
duricorius
050
045
055
052
053
058
041
050
039
060
067
044
044
042
027
050
072
058
065
078
039
065
057
071
065
052
052
052
062
059
067
044
072
074
056
056
047
056
065
Parasirocoiffaiti
044
045
055
054
055
058
056
047
050
052
079
050
041
060
056
080
051
064
061
086
042
064
060
062
058
052
056
057
068
053
052
081
063
068
056
056
055
053
064
Siro
acaroids
046
055
054
052
053
055
053
049
044
055
047
051
067
060
070
051
057
058
071
069
048
068
053
044
061
053
052
052
058
062
061
062
055
051
056
056
071
055
068
Siro
rubens
049
045
056
052
053
042
077
050
041
055
065
039
057
053
062
082
037
047
052
072
035
073
059
052
086
052
052
052
036
054
085
102
100
066
119
102
056
050
049
Suzukielus
sauteri
059
045
056
052
053
060
062
048
050
051
063
068
069
065
064
066
056
082
071
075
048
071
056
067
057
052
052
052
062
079
072
066
063
071
098
091
047
056
061
Stylocellusramblae
052
045
055
052
053
064
050
060
052
048
083
058
047
056
053
055
076
045
046
092
068
060
054
046
048
052
052
052
067
053
058
053
074
074
056
056
047
053
060
Stylocellustambusisi
074
045
057
052
053
066
049
059
046
067
058
052
050
041
029
059
076
047
056
082
066
060
057
070
056
052
052
052
061
047
055
041
065
074
056
056
047
052
062
Fan
gensiscavernarus
081
064
065
052
053
073
046
047
049
065
080
054
076
053
065
048
079
048
026
075
061
073
065
078
081
059
052
052
066
063
049
038
051
074
056
056
047
053
048
Fan
gensisspelaeus
073
045
057
052
053
078
093
047
050
061
093
065
077
056
064
076
053
068
030
076
054
069
056
069
066
052
052
052
076
068
055
086
061
074
056
056
047
057
054
374 Invertebrate Systematics B L de Bivort and G Giribet
4041
4243
4445
4647
4849
5051
5253
5455
5657
5859
6061
6263
6465
6667
6869
7071
7273
7475
7677
78
Parap
urcelliastaregaisp
nov
054
049
048
053
061
054
046
048
065
057
060
056
057
059
069
055
074
063
048
049
055
058
069
062
068
080
057
066
053
057
056
052
068
059
066
054
070
051
062
Parap
urcelliaam
atolaesp
nov
047
058
055
044
066
054
046
050
081
058
068
069
065
064
068
055
068
055
049
032
047
064
064
070
066
059
053
073
046
043
035
075
066
053
068
048
065
062
056
Parapurcelliaconvexa
spn
ov
036
030
068
095
053
054
055
050
064
057
053
052
056
065
070
055
069
066
055
062
053
069
052
075
070
059
063
067
049
055
069
062
077
034
062
057
071
052
062
Parapurcellianatalia
sp
nov
050
046
050
051
054
054
046
046
079
060
056
066
072
063
064
055
062
060
057
058
046
068
053
063
063
069
042
063
045
054
073
050
062
073
056
054
060
048
054
Parap
urcelliaminuta
spn
ov
046
046
054
041
030
054
047
049
075
060
059
050
058
059
066
055
062
062
057
040
062
066
072
061
062
076
044
064
052
056
058
076
095
069
053
058
062
057
054
Parap
urcelliaperegrinator
058
058
058
050
049
054
046
054
071
063
049
056
070
062
060
055
062
060
057
070
053
063
073
077
079
056
052
078
049
061
058
057
064
079
052
048
066
055
055
Parapurcelliasilvicola
052
054
062
050
044
054
050
048
064
058
063
059
055
062
069
055
064
061
052
051
062
067
072
070
066
019
045
064
059
059
048
061
074
066
065
073
070
064
057
Parapurcelliamonticola
067
073
086
060
055
054
053
051
066
057
050
062
071
063
070
055
060
065
058
053
044
064
061
039
072
075
047
080
048
061
056
047
066
061
066
047
061
053
056
Parap
urcelliarumpian
a040
035
072
081
058
054
055
045
075
057
048
067
074
063
065
055
082
059
056
072
052
066
051
064
073
068
049
076
073
066
056
058
070
035
054
090
078
093
058
Parapurcelliafissa
060
057
065
048
059
054
057
060
063
057
049
056
066
063
060
055
062
061
051
064
056
065
062
038
074
051
057
079
050
059
043
048
069
061
067
051
071
055
060
Purcelliatransvaalica
064
068
060
051
053
054
056
057
062
055
044
054
065
055
069
055
077
068
052
054
044
068
069
066
064
058
060
058
060
071
050
043
054
051
073
055
054
056
058
Purcelliaillustrans
111
080
101
074
037
054
082
067
066
052
047
046
060
066
057
055
056
056
055
047
052
066
063
057
066
048
055
077
052
068
051
043
059
048
053
047
072
054
059
Purcelliagriswoldisp
nov
067
078
063
048
049
054
054
049
065
057
045
050
066
061
065
055
053
061
059
046
050
063
074
056
070
071
070
066
058
070
035
050
051
069
049
058
077
052
056
Purcellialawrenceisp
nov
071
081
067
052
046
054
056
052
089
054
046
040
054
061
066
055
059
059
069
047
038
074
067
054
072
068
056
056
056
065
051
050
050
050
042
062
068
052
057
Speleosiro
argasiform
is057
056
054
045
060
054
047
044
055
066
041
058
078
061
045
055
058
061
057
044
076
049
090
079
055
054
085
058
077
089
050
052
051
070
043
041
050
048
060
Rakaiaisolate
066
075
061
060
062
054
061
079
032
073
059
069
071
063
059
055
066
060
066
044
061
052
039
053
069
056
038
058
049
050
049
077
071
058
077
062
065
064
059
Rakaialin
dsayi
050
060
058
056
068
054
057
054
070
057
067
065
060
067
060
055
055
056
068
059
073
059
060
044
078
074
049
053
060
059
050
064
057
085
059
055
064
066
058
Rakaiamag
naau
stralis
060
047
055
055
055
054
046
052
059
078
064
074
072
066
054
055
061
064
073
060
060
064
050
056
070
047
058
068
062
052
053
074
049
049
058
061
070
077
058
Rakaiamedia
060
082
041
052
062
054
061
052
065
078
064
082
075
061
058
055
060
062
075
060
047
059
043
059
063
071
060
064
049
037
076
056
070
050
061
063
070
051
060
Rakaiasolitaria
065
071
056
052
051
054
060
066
068
067
074
075
063
069
053
055
078
056
065
060
053
064
047
055
063
051
057
081
050
048
074
072
056
065
075
069
073
052
058
Rakaiasorensenidigitata
076
071
066
060
034
054
087
052
064
060
049
030
039
084
055
055
065
065
065
060
068
057
072
072
072
051
067
069
061
062
061
078
076
078
077
073
071
070
048
Aorakicrypta
071
069
053
060
054
054
082
048
068
067
068
063
051
069
055
055
059
073
071
054
064
047
075
078
071
056
080
040
068
049
067
081
042
064
068
041
047
057
048
Aorakidenticula
075
068
054
061
068
054
085
055
052
057
068
055
048
077
054
055
070
060
056
065
047
073
071
049
074
066
053
055
057
035
085
061
070
067
058
040
059
054
060
Aorakiinerma
069
070
071
078
065
054
069
046
068
060
064
063
052
072
050
055
081
064
054
050
058
060
062
072
042
055
057
073
070
068
054
052
047
043
079
046
057
055
061
Aorakilong
itarsa
069
066
057
062
073
054
067
078
064
059
075
063
056
071
056
055
065
073
057
069
073
043
074
071
064
050
071
054
064
049
071
078
050
052
062
056
063
057
062
Aorakitumidata
066
062
067
047
063
054
073
062
066
055
074
087
071
070
065
055
051
065
072
062
095
023
074
082
079
081
077
063
082
060
064
105
068
085
055
059
071
069
069
Karripurcellia
harveyi
050
061
072
074
086
054
084
076
057
054
047
047
064
062
057
055
056
054
061
077
075
053
060
073
053
047
066
059
064
070
054
051
055
061
045
052
054
050
060
Karripurcellia
peckorum
046
050
066
077
087
054
067
056
050
067
045
047
062
064
056
055
059
059
074
071
069
056
061
059
063
045
055
049
082
067
054
055
042
054
081
071
051
090
055
Karripurcellia
sierwaldae
050
052
064
067
068
054
062
061
060
050
060
044
044
064
065
055
052
060
051
060
060
058
057
064
061
060
063
070
079
074
051
040
045
058
056
081
057
097
048
Austropurcelliascoparia
079
078
069
066
053
054
074
064
051
052
080
056
037
089
038
055
056
057
080
050
057
076
067
062
060
048
053
056
052
062
064
060
061
068
062
066
062
052
063
Austrop
urcelliawoo
dwardi
072
067
074
061
054
054
075
079
049
062
075
067
047
074
036
055
058
058
053
054
051
086
061
052
056
059
038
054
054
065
060
046
058
049
063
080
067
054
048
Chileogovea
jocasta
067
064
065
054
054
054
061
052
054
055
069
066
052
069
045
055
049
058
057
057
063
066
042
053
059
068
068
059
056
074
054
056
057
060
043
057
046
050
048
Chileog
ovea
oedipu
s064
068
070
070
048
054
070
076
081
056
070
056
046
069
056
055
054
059
054
085
072
060
047
044
076
057
066
055
072
074
048
064
054
076
046
052
041
060
048
Pettaluslampetid
es063
055
059
058
058
054
057
051
063
063
053
077
084
052
075
055
058
051
057
058
054
052
051
057
060
063
062
052
073
067
054
051
039
075
083
040
034
053
061
Pettaluscimicifo
rmis
063
061
050
062
048
062
060
058
044
065
044
054
079
048
070
055
042
048
069
049
060
051
057
051
058
062
082
054
051
061
077
059
060
053
072
046
062
051
057
Troglosiroaelleni
059
056
029
089
066
085
058
054
066
046
078
071
050
062
045
090
056
060
068
076
055
055
066
058
045
064
076
054
064
065
059
049
055
055
046
049
037
052
060
Troglosirolongifo
ssa
059
055
058
070
077
085
077
053
061
045
069
071
058
048
054
093
038
065
070
085
059
056
052
082
044
051
071
049
056
045
075
053
050
053
038
063
073
064
048
Troglosironinq
ua066
051
072
073
079
083
077
061
060
041
068
067
053
045
064
092
062
059
058
067
059
053
070
075
052
061
082
049
075
076
046
055
048
051
036
053
045
068
048
Huitaca
ventralis
045
045
040
050
059
085
046
058
056
041
049
070
083
032
075
081
031
044
049
051
062
061
070
030
046
062
071
090
108
096
085
062
043
073
043
063
033
084
048
Neogo
veasp
042
067
051
086
073
087
071
066
031
046
039
066
087
037
059
055
066
130
104
061
066
071
062
039
042
072
075
076
074
077
079
055
050
036
084
064
038
095
061
Metag
ovea
philipi
057
068
061
073
063
085
062
044
040
041
066
069
059
039
057
055
061
044
080
064
056
085
058
045
035
059
041
075
059
063
053
049
046
047
065
054
034
059
048
Cypho
phthalmus
duricorius
060
046
053
056
061
086
050
058
065
056
053
064
073
054
051
087
060
064
057
082
060
061
060
065
048
047
064
036
055
052
070
059
066
065
071
077
072
064
078
Parasirocoiffaiti
056
064
071
050
066
081
046
074
063
060
089
070
051
051
055
078
055
065
053
062
063
060
032
073
042
079
053
052
044
048
078
068
088
061
060
077
070
055
048
Siro
acaroides
052
040
038
053
081
085
049
077
058
060
057
079
073
055
059
096
032
061
049
068
045
071
040
057
055
059
058
043
065
064
063
065
066
050
059
067
056
059
048
Siro
rubens
063
050
064
053
063
085
050
069
056
049
076
069
054
060
051
087
073
067
078
089
056
064
034
060
046
084
064
038
056
054
068
064
076
049
045
063
068
058
073
Suzukielus
sauteri
064
047
063
048
073
054
047
101
069
053
063
050
048
052
075
055
060
062
067
077
059
058
042
069
068
069
052
050
049
048
075
061
066
056
069
065
074
050
068
Stylocellusramblae
060
072
062
053
058
054
062
073
041
098
074
042
040
035
071
055
025
044
038
053
105
053
072
051
030
061
082
052
056
052
067
075
059
072
063
066
048
047
099
Stylocellustambusisi
056
063
061
052
060
054
056
069
047
096
073
050
041
040
067
055
063
044
038
042
064
060
074
050
036
048
063
044
060
049
067
060
058
078
057
085
046
062
087
Fan
gensiscavernarus
047
044
040
056
075
054
046
070
036
085
050
042
058
041
084
055
077
044
038
066
076
035
052
064
049
055
070
046
051
044
056
067
062
071
050
058
060
057
096
Fan
gensisspelaeus
049
061
044
055
079
054
046
070
047
086
046
035
058
042
105
055
087
044
038
065
077
029
068
065
060
084
050
047
051
045
078
070
083
057
054
071
065
056
096
Systematic revision of South African pettalids Invertebrate Systematics 375
combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)
Methods and abbreviations
Specimens used in this study were provided by the followinginstitutions
AMNH American Museum of Natural History New York(USA)
BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard
University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand
Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo
Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg
(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg
Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)
Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg
(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen
Copenhagen (Denmark)
Specimen handling and illustration
In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was
photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy
Character coding
Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3
Phylogenetic analyses
The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)
The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the
376 Invertebrate Systematics B L de Bivort and G Giribet
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
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idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
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Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
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Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
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pta
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cula
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ute
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ipu
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ren
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itata
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a s
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a p
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gn
a a
ust
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Pa
rap
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elli
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tare
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i sp
nov
Sir
o r
ub
en
s
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ro c
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i
Tro
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ata
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Pa
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elli
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Ne
og
ove
a s
p
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elli
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issa
Tro
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ifoss
a
Pa
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elli
a m
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la
Pe
tta
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lam
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Ao
raki
tu
mid
ata
Me
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a p
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pi
Au
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llia
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od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
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qu
a
Pa
rap
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elli
a s
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Ao
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erm
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Cyp
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uri
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us
Sp
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Ch
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us
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en
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Fa
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Fa
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Fa
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rum
Pu
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Ao
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tu
mid
ata
Pe
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lus
lam
pe
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es
Me
tag
ove
a p
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pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
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lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
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llia
sco
pa
ria
Ka
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elli
a p
eck
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m
Pa
rap
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elli
a a
ma
tola
sp
nov
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llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
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elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
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ov
Pa
rap
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elli
a s
tare
ga
i sp
nov
Pe
tta
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icifo
rmis
Pa
rap
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elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
The NMSA collections along with most collections ofpreviously described species were made before the era ofmolecular phylogenetics Their long-term storage in 70ethanol has precluded the possibility of analysing the species-level relationships of the South African pettalids using molecular
sequences In order to investigate the phylogenetic relationshipsof the SouthAfricanmaterial we chose to perform aphylogeneticanalysis under the parsimony criterion using all availablemorphological data Attempts at lsquototal evidencersquo analysis haveproved successful in past analyses of the Cyphophthalmi
Table 1 Discrete character matrixCharacter codings are explained in Appendix 2
Taxa Characters000000000 1111111111 2222222222 3333333333 4444444444 5555555555 666
123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012
Parapurcellia staregai sp nov 032200011 031101100 011001100 00-000100 0112000100 1130100100 0
Parapurcellia amatolae sp nov 032200011 031100100 1101011000 00-0020100 0112000100 1130100100 0
Parapurcellia convexa sp nov 032200011 031100100 0101011100 00-0020100 0112000110 1120100101 0
Parapurcellia natalia sp nov 032200011 031000100 011101100 00-000100 0112000110 1130100100 0
Parapurcellia minuta sp nov 032200011 031101100 0101011000 00-0020110 0112000110 10-0100100 0
Parapurcellia peregrinator 032200011 021100100 111101110 00-00300 0112000110 1120100100 0
Parapurcellia silvicola 032200011 031100000 0101011100 00-0020100 01100110 0100100101 0
Parapurcellia monticola 032200011 021100000 0101011100 00-0020100 01100110 1130100100 0
Parapurcellia rumpiana 032200011 021001100 010111101 0-30100 01100110 110100100 0
Parapurcellia fissa 032200011 031100100 0101110 30100 0100110 130100100 0
Purcellia transvaalica 222200011 020101100 110101100 00-0020 0112100100 1140110100 0
Purcellia illustrans 222200011 000100000 1101011101 00-0020100 0112100100 0140110100 001
Purcellia griswoldi sp nov 222200010 020100100 1001011100 00-0020100 0112100100 1140110100 011
Purcellia lawrencei sp nov 222200011 030100100 110101100 00-0020100 0112100100 1140110100 0
Speleosiro argasiformis 022200011 000100000 1100011101 0-0020100 0111000100 0140110100 0
Rakaia isolata 232200011 000101000 110101100 00-0020000 0100110 0100101101 0
Rakaia lindsayi 232200011 000100000 1100010210 00-0020000 01100110 0100100101 0
Rakaia magna australis 232200011 000101000 1100011200 00-0020000 01100110 0100100100 0
Rakaia media 232200011 000100000 1101011200 00-0020000 011200110 0100100100 0
Rakaia solitaria 132200011 000100000 1101011200 00-0020000 01100110 0100101100 0
Rakaia sorenseni digitata 232200011 000100000 1101011100 00-0020050 010ndashndash00110 00-01-1000 0
Aoraki crypta 232200011 000100000 1110010101 00-0020000 011200110 01311-0000 0
Aoraki denticula 232200011 000100000 1110010201 00-0021000 011200110 01301-0001 0
Aoraki inerma 232200011 000100000 1110010201 00-0020000 011200110 01311-0000 0
Aoraki longitarsa 232200011 000100000 1110010201 00-0021000 0100110 0100100101 1
Aoraki tumidata 232200011 000101000 11001011 0-2 112100111 10-0101101 0
Karripurcellia harveyi 232200011 000100010 1110010210 00-0020000 010ndashndash10100 00-00-0000 0
Karripurcellia peckorum 232200011 000101010 1110010210 00-0020000 010ndashndash10100 00-00-0000 001
Karripurcellia sierwaldae 232200011 000101010 1100011200 00-0020000 010ndashndash10100 00-00-0000 0
Austropurcellia scoparia 232200011 000100000 1101011101 00-0030100 0114000110 01101-0000 11
Austropurcellia woodwardi 232200011 020100000 111101 00-0030100 011000110 0130100100 1
Chileogovea jocasta 132200011 001100000 1110010101 00-1140000 0112100100 00-00-0000 0
Chileogovea oedipus 132100011 001100000 1110010211 00-1141000 0112100101 10-00-0000 01
Pettalus lampetides 122200011 001100010 1110110001 00-0041001 010ndashndash02100 00-01-1000 01
Pettalus cimiciformis 122100011 00310001 1110010101 00-4100 010ndashndash02100 00-01-1000 0
Troglosiro aelleni 010210011 001000000 2110000201 0110030000 001003-00 00-00-0000 0
Troglosiro longifossa 010210011 0010100100 2110000101 0110030000 001003-00 00-00-0010 0
Troglosiro ninqua 010210011 0010100000 2110000000 0110030000 001003-00 00-00-0010 0
Huitaca ventralis 010200101 111-100010 2110010201 0110030001 001003-00 00-00-0000 00
Neogovea sp 010200111 1310101010 1101000 0 003-00 00-00-0000 0
Metagovea philipi 010200111 12110000 2110000101 0110030001 001003-00 00-00-0000 001
Cyphophthalmus duricorius 010211011 0020000101 0000010100 00-0010000 001103-01 10-00-0000 011
Parasiro coiffaiti 000201010 0010000101 0100010000 0111120000 000ndashndash01000 00-00-0000 010
Siro acaroides 010211010 0010000100 0000000000 00-1110000 0011103-00 10-01-0000 011
Siro rubens 010211011 0020000100 0000020100 00-0010000 0011103-01 00-00-0000 011
Suzukielus sauteri 012200011 001000100 0000020100 1101120100 0013000101 00-00-0000 011
Stylocellus ramblae 111200101 000010001 011011021 00-0151010 020ndashndash00000 00-00-0000 0
Stylocellus tambusisi 111200100 121010001 01100102 00-001010 020ndashndash00000 10-00-0000 0
Fangensis cavernarus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001
Fangensis spelaeus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001
Systematic revision of South African pettalids Invertebrate Systematics 373
Tab
le2
Con
tinu
ouscharactermatrixas
itap
pearsin
the11
weigh
tedcombinedan
alysis
Character
definitio
nsaregivenin
App
endix3
12
34
56
78
910
1112
1314
1516
1718
1920
2122
2324
2526
2728
2930
3132
3334
3536
3738
39
Parap
urcelliastaregaisp
nov
054
059
058
052
053
053
059
047
064
051
058
063
047
067
069
075
048
071
070
056
055
049
062
073
057
057
052
052
054
066
054
068
048
048
056
056
067
061
052
Parap
urcelliaam
atolaesp
nov
047
061
057
056
056
064
052
047
065
057
053
072
051
062
061
059
062
070
066
056
042
063
062
053
053
058
058
057
072
055
050
061
043
049
056
056
066
062
049
Parapurcelliaconvexa
spn
ov
048
069
058
075
067
066
049
053
060
051
050
078
081
063
074
051
064
062
071
059
056
065
057
059
045
066
085
075
071
070
068
052
057
051
056
056
061
060
054
Parapurcellianatalia
sp
nov
044
056
057
052
053
057
049
047
057
049
060
063
048
050
071
061
105
082
074
060
053
059
061
076
059
059
052
052
065
056
043
065
043
047
056
056
071
058
049
Parapurcelliaminuta
spn
ov
042
056
056
052
053
046
050
047
057
054
081
075
047
068
072
082
048
069
062
060
059
069
059
050
052
057
052
052
043
067
055
064
057
052
056
056
067
058
048
Parapurcelliaperegrinator
059
061
058
059
055
057
059
047
077
055
065
065
048
063
049
048
066
061
057
061
055
056
057
062
071
059
060
055
058
061
053
062
053
047
056
056
065
059
059
Parapurcelliasilvicola
055
063
063
076
064
049
062
050
062
062
060
072
077
053
079
050
065
079
070
059
060
065
064
064
057
071
080
067
047
097
059
068
040
059
056
056
064
058
070
Parapurcelliamonticola
054
061
059
061
053
056
045
053
073
052
058
074
045
055
064
073
050
070
059
057
065
062
058
064
061
063
063
052
058
054
066
049
074
051
056
056
066
063
062
Parap
urcelliarumpiana
051
059
056
057
053
050
040
053
067
078
046
073
075
057
073
080
037
074
071
057
061
058
054
056
059
059
060
052
049
062
066
039
061
046
056
056
075
060
055
Parapurcelliafissa
043
064
064
052
053
056
059
053
065
052
055
070
071
069
076
063
055
066
060
057
063
079
078
060
051
085
052
052
062
036
068
083
053
053
056
056
068
060
052
Purcelliatransvaalica
069
064
057
052
053
048
060
058
061
058
041
071
070
057
074
050
061
053
072
050
067
050
056
054
056
056
052
052
046
056
054
051
050
036
056
056
074
058
048
Purcelliaillustrans
072
063
055
052
053
052
060
087
081
056
061
063
062
061
077
049
059
075
071
051
065
058
051
078
087
054
052
052
050
089
096
053
076
036
056
056
078
061
042
Purcelliagriswoldisp
nov
062
063
057
057
053
043
062
055
067
053
065
066
062
044
064
068
050
057
058
048
075
059
055
053
058
058
057
052
041
061
061
060
060
040
056
056
075
057
045
Purcellialawrenceisp
nov
062
061
056
057
053
051
066
057
057
053
059
069
067
047
054
066
053
068
067
048
080
063
052
056
050
055
057
052
050
063
068
064
068
039
056
056
075
053
044
Speleosiro
argasiform
is097
077
060
075
059
073
091
048
071
051
049
044
063
039
056
055
068
031
035
045
108
046
054
047
050
057
064
056
061
053
056
060
057
047
056
056
070
058
043
Rakaiaisolate
059
077
067
102
058
069
066
061
059
063
067
077
077
070
062
046
077
054
068
052
062
062
072
061
043
076
104
058
071
066
061
068
059
062
056
056
058
064
099
Rakaialin
dsayi
061
063
062
083
053
059
053
057
054
066
071
056
071
077
067
086
046
079
059
051
057
071
058
057
061
070
082
052
061
063
037
055
037
056
056
056
062
066
062
Rakaiamag
naau
stralis
074
068
064
078
061
085
070
047
077
085
072
052
071
057
043
053
067
068
062
054
062
062
066
068
058
060
072
059
078
065
065
059
067
053
056
056
065
058
075
Rakaiamedia
062
070
062
063
057
055
077
062
066
052
066
058
070
081
066
087
041
065
064
049
057
057
068
058
068
061
062
057
054
058
054
077
057
064
056
056
061
063
077
Rakaiasolitaria
057
065
059
071
053
065
052
060
079
053
074
080
051
055
051
046
070
065
068
053
077
043
055
037
060
065
073
052
069
069
067
055
065
066
056
056
053
062
074
Rakaiasorensenidigitata
057
071
055
054
055
075
071
080
065
066
056
093
052
085
038
054
064
069
065
051
075
075
053
066
069
054
055
056
083
060
059
080
058
074
056
056
047
053
078
Aorakicrypta
071
079
061
058
059
085
069
088
041
059
049
059
043
072
059
047
071
060
064
052
060
037
051
039
053
071
057
058
081
067
062
061
053
039
056
056
077
075
086
Aorakidenticula
057
067
055
068
062
063
048
080
054
054
056
065
051
085
058
060
067
068
071
053
066
065
051
057
069
055
071
064
066
059
063
050
063
044
075
072
071
070
070
Aorakiinerma
067
066
057
063
063
062
056
070
075
056
044
066
065
077
053
051
066
060
059
054
073
066
053
053
063
057
062
063
060
054
060
050
059
062
056
056
059
072
074
Aorakilong
itarsa
066
076
073
098
109
065
058
069
056
058
083
056
049
062
028
049
061
063
064
050
061
059
066
079
060
091
094
107
064
076
062
055
058
064
056
056
059
129
062
Aorakitumidata
064
072
058
081
069
079
070
072
063
068
071
073
079
083
054
080
060
073
070
050
062
072
055
054
062
060
083
070
081
085
062
070
050
070
075
096
080
059
071
Karripurcellia
harveyi
067
052
058
061
066
051
079
089
062
052
069
056
073
060
074
075
037
050
043
055
056
063
060
069
091
052
060
065
049
066
050
072
048
067
056
056
047
057
058
Karripurcellia
peckorum
068
062
057
071
068
069
062
070
051
094
057
064
070
057
052
045
071
046
037
055
060
064
057
059
076
054
069
067
064
070
046
053
037
068
056
056
047
056
056
Karripurcellia
sierwaldae
056
056
057
052
053
042
062
061
067
088
056
061
076
063
070
057
060
043
032
055
062
069
062
067
068
053
052
052
039
067
057
064
051
073
064
087
047
057
060
Austropurcelliascoparia
055
050
054
052
053
067
052
068
054
051
045
065
050
064
057
044
071
065
074
059
071
072
051
054
084
053
052
052
071
052
082
055
088
062
056
056
061
091
053
Austrop
urcelliawoo
dwardi
054
065
058
052
058
056
061
069
053
053
042
066
049
066
068
053
057
062
068
054
063
047
049
036
050
067
052
060
056
065
079
064
068
059
056
056
047
061
053
Chileogovea
jocasta
049
053
055
052
054
058
050
057
055
049
047
049
058
060
055
063
050
050
058
051
074
055
053
053
060
053
052
055
061
057
069
057
068
066
082
088
082
053
069
Chileog
ovea
Oedipus
070
053
054
052
053
064
070
074
057
062
053
052
043
064
054
051
058
072
078
053
071
012
049
043
054
052
052
052
061
052
063
061
070
062
092
091
083
051
048
Pettaluslampetid
es063
066
078
052
092
067
057
058
053
055
063
039
048
055
057
063
064
068
068
046
064
050
095
090
044
084
052
093
065
042
050
052
066
074
056
056
047
052
055
Pettaluscimicifo
rmis
100
110
140
066
095
089
082
076
066
058
074
033
039
021
057
058
058
056
046
041
056
085
125
064
044
116
059
074
070
039
047
055
071
068
056
056
047
050
061
Troglosiroaelleni
059
053
059
054
066
066
064
057
054
053
052
040
067
066
072
060
064
044
053
076
045
064
066
075
062
054
054
067
068
054
061
066
064
074
056
056
047
055
058
Troglosirolong
ifossa
051
045
055
052
073
064
048
070
049
060
057
042
057
053
069
043
069
053
064
079
041
059
055
061
066
052
052
081
070
049
049
052
055
074
056
056
047
056
053
Troglosironinq
ua059
045
057
053
086
068
049
074
049
066
048
044
074
054
064
052
063
048
059
076
044
054
063
072
068
052
054
092
069
062
076
048
070
074
056
056
047
057
072
Huitaca
ventralis
075
045
057
052
060
036
084
047
105
094
063
054
058
067
072
068
054
040
056
075
057
046
058
070
053
052
052
058
039
045
038
057
047
074
056
056
047
052
048
Neogo
veasp
067
056
058
065
070
031
070
075
081
099
042
055
067
047
063
068
042
031
045
074
050
038
061
073
061
052
063
069
033
035
035
055
044
074
056
056
047
050
045
Metagovea
philipi
045
045
054
052
053
041
045
057
065
055
056
056
066
051
053
054
056
048
062
078
057
057
053
050
031
052
052
052
036
042
056
048
073
074
056
056
047
053
068
Cypho
phthalmus
duricorius
050
045
055
052
053
058
041
050
039
060
067
044
044
042
027
050
072
058
065
078
039
065
057
071
065
052
052
052
062
059
067
044
072
074
056
056
047
056
065
Parasirocoiffaiti
044
045
055
054
055
058
056
047
050
052
079
050
041
060
056
080
051
064
061
086
042
064
060
062
058
052
056
057
068
053
052
081
063
068
056
056
055
053
064
Siro
acaroids
046
055
054
052
053
055
053
049
044
055
047
051
067
060
070
051
057
058
071
069
048
068
053
044
061
053
052
052
058
062
061
062
055
051
056
056
071
055
068
Siro
rubens
049
045
056
052
053
042
077
050
041
055
065
039
057
053
062
082
037
047
052
072
035
073
059
052
086
052
052
052
036
054
085
102
100
066
119
102
056
050
049
Suzukielus
sauteri
059
045
056
052
053
060
062
048
050
051
063
068
069
065
064
066
056
082
071
075
048
071
056
067
057
052
052
052
062
079
072
066
063
071
098
091
047
056
061
Stylocellusramblae
052
045
055
052
053
064
050
060
052
048
083
058
047
056
053
055
076
045
046
092
068
060
054
046
048
052
052
052
067
053
058
053
074
074
056
056
047
053
060
Stylocellustambusisi
074
045
057
052
053
066
049
059
046
067
058
052
050
041
029
059
076
047
056
082
066
060
057
070
056
052
052
052
061
047
055
041
065
074
056
056
047
052
062
Fan
gensiscavernarus
081
064
065
052
053
073
046
047
049
065
080
054
076
053
065
048
079
048
026
075
061
073
065
078
081
059
052
052
066
063
049
038
051
074
056
056
047
053
048
Fan
gensisspelaeus
073
045
057
052
053
078
093
047
050
061
093
065
077
056
064
076
053
068
030
076
054
069
056
069
066
052
052
052
076
068
055
086
061
074
056
056
047
057
054
374 Invertebrate Systematics B L de Bivort and G Giribet
4041
4243
4445
4647
4849
5051
5253
5455
5657
5859
6061
6263
6465
6667
6869
7071
7273
7475
7677
78
Parap
urcelliastaregaisp
nov
054
049
048
053
061
054
046
048
065
057
060
056
057
059
069
055
074
063
048
049
055
058
069
062
068
080
057
066
053
057
056
052
068
059
066
054
070
051
062
Parap
urcelliaam
atolaesp
nov
047
058
055
044
066
054
046
050
081
058
068
069
065
064
068
055
068
055
049
032
047
064
064
070
066
059
053
073
046
043
035
075
066
053
068
048
065
062
056
Parapurcelliaconvexa
spn
ov
036
030
068
095
053
054
055
050
064
057
053
052
056
065
070
055
069
066
055
062
053
069
052
075
070
059
063
067
049
055
069
062
077
034
062
057
071
052
062
Parapurcellianatalia
sp
nov
050
046
050
051
054
054
046
046
079
060
056
066
072
063
064
055
062
060
057
058
046
068
053
063
063
069
042
063
045
054
073
050
062
073
056
054
060
048
054
Parap
urcelliaminuta
spn
ov
046
046
054
041
030
054
047
049
075
060
059
050
058
059
066
055
062
062
057
040
062
066
072
061
062
076
044
064
052
056
058
076
095
069
053
058
062
057
054
Parap
urcelliaperegrinator
058
058
058
050
049
054
046
054
071
063
049
056
070
062
060
055
062
060
057
070
053
063
073
077
079
056
052
078
049
061
058
057
064
079
052
048
066
055
055
Parapurcelliasilvicola
052
054
062
050
044
054
050
048
064
058
063
059
055
062
069
055
064
061
052
051
062
067
072
070
066
019
045
064
059
059
048
061
074
066
065
073
070
064
057
Parapurcelliamonticola
067
073
086
060
055
054
053
051
066
057
050
062
071
063
070
055
060
065
058
053
044
064
061
039
072
075
047
080
048
061
056
047
066
061
066
047
061
053
056
Parap
urcelliarumpian
a040
035
072
081
058
054
055
045
075
057
048
067
074
063
065
055
082
059
056
072
052
066
051
064
073
068
049
076
073
066
056
058
070
035
054
090
078
093
058
Parapurcelliafissa
060
057
065
048
059
054
057
060
063
057
049
056
066
063
060
055
062
061
051
064
056
065
062
038
074
051
057
079
050
059
043
048
069
061
067
051
071
055
060
Purcelliatransvaalica
064
068
060
051
053
054
056
057
062
055
044
054
065
055
069
055
077
068
052
054
044
068
069
066
064
058
060
058
060
071
050
043
054
051
073
055
054
056
058
Purcelliaillustrans
111
080
101
074
037
054
082
067
066
052
047
046
060
066
057
055
056
056
055
047
052
066
063
057
066
048
055
077
052
068
051
043
059
048
053
047
072
054
059
Purcelliagriswoldisp
nov
067
078
063
048
049
054
054
049
065
057
045
050
066
061
065
055
053
061
059
046
050
063
074
056
070
071
070
066
058
070
035
050
051
069
049
058
077
052
056
Purcellialawrenceisp
nov
071
081
067
052
046
054
056
052
089
054
046
040
054
061
066
055
059
059
069
047
038
074
067
054
072
068
056
056
056
065
051
050
050
050
042
062
068
052
057
Speleosiro
argasiform
is057
056
054
045
060
054
047
044
055
066
041
058
078
061
045
055
058
061
057
044
076
049
090
079
055
054
085
058
077
089
050
052
051
070
043
041
050
048
060
Rakaiaisolate
066
075
061
060
062
054
061
079
032
073
059
069
071
063
059
055
066
060
066
044
061
052
039
053
069
056
038
058
049
050
049
077
071
058
077
062
065
064
059
Rakaialin
dsayi
050
060
058
056
068
054
057
054
070
057
067
065
060
067
060
055
055
056
068
059
073
059
060
044
078
074
049
053
060
059
050
064
057
085
059
055
064
066
058
Rakaiamag
naau
stralis
060
047
055
055
055
054
046
052
059
078
064
074
072
066
054
055
061
064
073
060
060
064
050
056
070
047
058
068
062
052
053
074
049
049
058
061
070
077
058
Rakaiamedia
060
082
041
052
062
054
061
052
065
078
064
082
075
061
058
055
060
062
075
060
047
059
043
059
063
071
060
064
049
037
076
056
070
050
061
063
070
051
060
Rakaiasolitaria
065
071
056
052
051
054
060
066
068
067
074
075
063
069
053
055
078
056
065
060
053
064
047
055
063
051
057
081
050
048
074
072
056
065
075
069
073
052
058
Rakaiasorensenidigitata
076
071
066
060
034
054
087
052
064
060
049
030
039
084
055
055
065
065
065
060
068
057
072
072
072
051
067
069
061
062
061
078
076
078
077
073
071
070
048
Aorakicrypta
071
069
053
060
054
054
082
048
068
067
068
063
051
069
055
055
059
073
071
054
064
047
075
078
071
056
080
040
068
049
067
081
042
064
068
041
047
057
048
Aorakidenticula
075
068
054
061
068
054
085
055
052
057
068
055
048
077
054
055
070
060
056
065
047
073
071
049
074
066
053
055
057
035
085
061
070
067
058
040
059
054
060
Aorakiinerma
069
070
071
078
065
054
069
046
068
060
064
063
052
072
050
055
081
064
054
050
058
060
062
072
042
055
057
073
070
068
054
052
047
043
079
046
057
055
061
Aorakilong
itarsa
069
066
057
062
073
054
067
078
064
059
075
063
056
071
056
055
065
073
057
069
073
043
074
071
064
050
071
054
064
049
071
078
050
052
062
056
063
057
062
Aorakitumidata
066
062
067
047
063
054
073
062
066
055
074
087
071
070
065
055
051
065
072
062
095
023
074
082
079
081
077
063
082
060
064
105
068
085
055
059
071
069
069
Karripurcellia
harveyi
050
061
072
074
086
054
084
076
057
054
047
047
064
062
057
055
056
054
061
077
075
053
060
073
053
047
066
059
064
070
054
051
055
061
045
052
054
050
060
Karripurcellia
peckorum
046
050
066
077
087
054
067
056
050
067
045
047
062
064
056
055
059
059
074
071
069
056
061
059
063
045
055
049
082
067
054
055
042
054
081
071
051
090
055
Karripurcellia
sierwaldae
050
052
064
067
068
054
062
061
060
050
060
044
044
064
065
055
052
060
051
060
060
058
057
064
061
060
063
070
079
074
051
040
045
058
056
081
057
097
048
Austropurcelliascoparia
079
078
069
066
053
054
074
064
051
052
080
056
037
089
038
055
056
057
080
050
057
076
067
062
060
048
053
056
052
062
064
060
061
068
062
066
062
052
063
Austrop
urcelliawoo
dwardi
072
067
074
061
054
054
075
079
049
062
075
067
047
074
036
055
058
058
053
054
051
086
061
052
056
059
038
054
054
065
060
046
058
049
063
080
067
054
048
Chileogovea
jocasta
067
064
065
054
054
054
061
052
054
055
069
066
052
069
045
055
049
058
057
057
063
066
042
053
059
068
068
059
056
074
054
056
057
060
043
057
046
050
048
Chileog
ovea
oedipu
s064
068
070
070
048
054
070
076
081
056
070
056
046
069
056
055
054
059
054
085
072
060
047
044
076
057
066
055
072
074
048
064
054
076
046
052
041
060
048
Pettaluslampetid
es063
055
059
058
058
054
057
051
063
063
053
077
084
052
075
055
058
051
057
058
054
052
051
057
060
063
062
052
073
067
054
051
039
075
083
040
034
053
061
Pettaluscimicifo
rmis
063
061
050
062
048
062
060
058
044
065
044
054
079
048
070
055
042
048
069
049
060
051
057
051
058
062
082
054
051
061
077
059
060
053
072
046
062
051
057
Troglosiroaelleni
059
056
029
089
066
085
058
054
066
046
078
071
050
062
045
090
056
060
068
076
055
055
066
058
045
064
076
054
064
065
059
049
055
055
046
049
037
052
060
Troglosirolongifo
ssa
059
055
058
070
077
085
077
053
061
045
069
071
058
048
054
093
038
065
070
085
059
056
052
082
044
051
071
049
056
045
075
053
050
053
038
063
073
064
048
Troglosironinq
ua066
051
072
073
079
083
077
061
060
041
068
067
053
045
064
092
062
059
058
067
059
053
070
075
052
061
082
049
075
076
046
055
048
051
036
053
045
068
048
Huitaca
ventralis
045
045
040
050
059
085
046
058
056
041
049
070
083
032
075
081
031
044
049
051
062
061
070
030
046
062
071
090
108
096
085
062
043
073
043
063
033
084
048
Neogo
veasp
042
067
051
086
073
087
071
066
031
046
039
066
087
037
059
055
066
130
104
061
066
071
062
039
042
072
075
076
074
077
079
055
050
036
084
064
038
095
061
Metag
ovea
philipi
057
068
061
073
063
085
062
044
040
041
066
069
059
039
057
055
061
044
080
064
056
085
058
045
035
059
041
075
059
063
053
049
046
047
065
054
034
059
048
Cypho
phthalmus
duricorius
060
046
053
056
061
086
050
058
065
056
053
064
073
054
051
087
060
064
057
082
060
061
060
065
048
047
064
036
055
052
070
059
066
065
071
077
072
064
078
Parasirocoiffaiti
056
064
071
050
066
081
046
074
063
060
089
070
051
051
055
078
055
065
053
062
063
060
032
073
042
079
053
052
044
048
078
068
088
061
060
077
070
055
048
Siro
acaroides
052
040
038
053
081
085
049
077
058
060
057
079
073
055
059
096
032
061
049
068
045
071
040
057
055
059
058
043
065
064
063
065
066
050
059
067
056
059
048
Siro
rubens
063
050
064
053
063
085
050
069
056
049
076
069
054
060
051
087
073
067
078
089
056
064
034
060
046
084
064
038
056
054
068
064
076
049
045
063
068
058
073
Suzukielus
sauteri
064
047
063
048
073
054
047
101
069
053
063
050
048
052
075
055
060
062
067
077
059
058
042
069
068
069
052
050
049
048
075
061
066
056
069
065
074
050
068
Stylocellusramblae
060
072
062
053
058
054
062
073
041
098
074
042
040
035
071
055
025
044
038
053
105
053
072
051
030
061
082
052
056
052
067
075
059
072
063
066
048
047
099
Stylocellustambusisi
056
063
061
052
060
054
056
069
047
096
073
050
041
040
067
055
063
044
038
042
064
060
074
050
036
048
063
044
060
049
067
060
058
078
057
085
046
062
087
Fan
gensiscavernarus
047
044
040
056
075
054
046
070
036
085
050
042
058
041
084
055
077
044
038
066
076
035
052
064
049
055
070
046
051
044
056
067
062
071
050
058
060
057
096
Fan
gensisspelaeus
049
061
044
055
079
054
046
070
047
086
046
035
058
042
105
055
087
044
038
065
077
029
068
065
060
084
050
047
051
045
078
070
083
057
054
071
065
056
096
Systematic revision of South African pettalids Invertebrate Systematics 375
combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)
Methods and abbreviations
Specimens used in this study were provided by the followinginstitutions
AMNH American Museum of Natural History New York(USA)
BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard
University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand
Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo
Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg
(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg
Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)
Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg
(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen
Copenhagen (Denmark)
Specimen handling and illustration
In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was
photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy
Character coding
Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3
Phylogenetic analyses
The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)
The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the
376 Invertebrate Systematics B L de Bivort and G Giribet
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
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trict
consensusof
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underparsim
onyanalysisof
thecontinuous
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pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
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phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
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81
41
21
11
12
14
18
Pet
talid
ae S
tylo
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Neo
gove
idae
Tro
glos
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(T
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Neo
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)K
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trop
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ogov
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(Pur
celli
a gr
isw
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renc
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p n
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(Pet
talid
ae +
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ukie
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(Aor
aki +
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(Pet
talid
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nida
e)
(Kar
ripur
celli
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talu
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(Par
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celli
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leos
iro)
(Pur
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Aus
trop
urce
llia)
AB
C
Ao
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cula
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Tro
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a Ka
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arv
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99 51 69
75 92 95 8581
80 69
94
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74 83
86 86 5665
6379
8877 100
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81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Tab
le2
Con
tinu
ouscharactermatrixas
itap
pearsin
the11
weigh
tedcombinedan
alysis
Character
definitio
nsaregivenin
App
endix3
12
34
56
78
910
1112
1314
1516
1718
1920
2122
2324
2526
2728
2930
3132
3334
3536
3738
39
Parap
urcelliastaregaisp
nov
054
059
058
052
053
053
059
047
064
051
058
063
047
067
069
075
048
071
070
056
055
049
062
073
057
057
052
052
054
066
054
068
048
048
056
056
067
061
052
Parap
urcelliaam
atolaesp
nov
047
061
057
056
056
064
052
047
065
057
053
072
051
062
061
059
062
070
066
056
042
063
062
053
053
058
058
057
072
055
050
061
043
049
056
056
066
062
049
Parapurcelliaconvexa
spn
ov
048
069
058
075
067
066
049
053
060
051
050
078
081
063
074
051
064
062
071
059
056
065
057
059
045
066
085
075
071
070
068
052
057
051
056
056
061
060
054
Parapurcellianatalia
sp
nov
044
056
057
052
053
057
049
047
057
049
060
063
048
050
071
061
105
082
074
060
053
059
061
076
059
059
052
052
065
056
043
065
043
047
056
056
071
058
049
Parapurcelliaminuta
spn
ov
042
056
056
052
053
046
050
047
057
054
081
075
047
068
072
082
048
069
062
060
059
069
059
050
052
057
052
052
043
067
055
064
057
052
056
056
067
058
048
Parapurcelliaperegrinator
059
061
058
059
055
057
059
047
077
055
065
065
048
063
049
048
066
061
057
061
055
056
057
062
071
059
060
055
058
061
053
062
053
047
056
056
065
059
059
Parapurcelliasilvicola
055
063
063
076
064
049
062
050
062
062
060
072
077
053
079
050
065
079
070
059
060
065
064
064
057
071
080
067
047
097
059
068
040
059
056
056
064
058
070
Parapurcelliamonticola
054
061
059
061
053
056
045
053
073
052
058
074
045
055
064
073
050
070
059
057
065
062
058
064
061
063
063
052
058
054
066
049
074
051
056
056
066
063
062
Parap
urcelliarumpiana
051
059
056
057
053
050
040
053
067
078
046
073
075
057
073
080
037
074
071
057
061
058
054
056
059
059
060
052
049
062
066
039
061
046
056
056
075
060
055
Parapurcelliafissa
043
064
064
052
053
056
059
053
065
052
055
070
071
069
076
063
055
066
060
057
063
079
078
060
051
085
052
052
062
036
068
083
053
053
056
056
068
060
052
Purcelliatransvaalica
069
064
057
052
053
048
060
058
061
058
041
071
070
057
074
050
061
053
072
050
067
050
056
054
056
056
052
052
046
056
054
051
050
036
056
056
074
058
048
Purcelliaillustrans
072
063
055
052
053
052
060
087
081
056
061
063
062
061
077
049
059
075
071
051
065
058
051
078
087
054
052
052
050
089
096
053
076
036
056
056
078
061
042
Purcelliagriswoldisp
nov
062
063
057
057
053
043
062
055
067
053
065
066
062
044
064
068
050
057
058
048
075
059
055
053
058
058
057
052
041
061
061
060
060
040
056
056
075
057
045
Purcellialawrenceisp
nov
062
061
056
057
053
051
066
057
057
053
059
069
067
047
054
066
053
068
067
048
080
063
052
056
050
055
057
052
050
063
068
064
068
039
056
056
075
053
044
Speleosiro
argasiform
is097
077
060
075
059
073
091
048
071
051
049
044
063
039
056
055
068
031
035
045
108
046
054
047
050
057
064
056
061
053
056
060
057
047
056
056
070
058
043
Rakaiaisolate
059
077
067
102
058
069
066
061
059
063
067
077
077
070
062
046
077
054
068
052
062
062
072
061
043
076
104
058
071
066
061
068
059
062
056
056
058
064
099
Rakaialin
dsayi
061
063
062
083
053
059
053
057
054
066
071
056
071
077
067
086
046
079
059
051
057
071
058
057
061
070
082
052
061
063
037
055
037
056
056
056
062
066
062
Rakaiamag
naau
stralis
074
068
064
078
061
085
070
047
077
085
072
052
071
057
043
053
067
068
062
054
062
062
066
068
058
060
072
059
078
065
065
059
067
053
056
056
065
058
075
Rakaiamedia
062
070
062
063
057
055
077
062
066
052
066
058
070
081
066
087
041
065
064
049
057
057
068
058
068
061
062
057
054
058
054
077
057
064
056
056
061
063
077
Rakaiasolitaria
057
065
059
071
053
065
052
060
079
053
074
080
051
055
051
046
070
065
068
053
077
043
055
037
060
065
073
052
069
069
067
055
065
066
056
056
053
062
074
Rakaiasorensenidigitata
057
071
055
054
055
075
071
080
065
066
056
093
052
085
038
054
064
069
065
051
075
075
053
066
069
054
055
056
083
060
059
080
058
074
056
056
047
053
078
Aorakicrypta
071
079
061
058
059
085
069
088
041
059
049
059
043
072
059
047
071
060
064
052
060
037
051
039
053
071
057
058
081
067
062
061
053
039
056
056
077
075
086
Aorakidenticula
057
067
055
068
062
063
048
080
054
054
056
065
051
085
058
060
067
068
071
053
066
065
051
057
069
055
071
064
066
059
063
050
063
044
075
072
071
070
070
Aorakiinerma
067
066
057
063
063
062
056
070
075
056
044
066
065
077
053
051
066
060
059
054
073
066
053
053
063
057
062
063
060
054
060
050
059
062
056
056
059
072
074
Aorakilong
itarsa
066
076
073
098
109
065
058
069
056
058
083
056
049
062
028
049
061
063
064
050
061
059
066
079
060
091
094
107
064
076
062
055
058
064
056
056
059
129
062
Aorakitumidata
064
072
058
081
069
079
070
072
063
068
071
073
079
083
054
080
060
073
070
050
062
072
055
054
062
060
083
070
081
085
062
070
050
070
075
096
080
059
071
Karripurcellia
harveyi
067
052
058
061
066
051
079
089
062
052
069
056
073
060
074
075
037
050
043
055
056
063
060
069
091
052
060
065
049
066
050
072
048
067
056
056
047
057
058
Karripurcellia
peckorum
068
062
057
071
068
069
062
070
051
094
057
064
070
057
052
045
071
046
037
055
060
064
057
059
076
054
069
067
064
070
046
053
037
068
056
056
047
056
056
Karripurcellia
sierwaldae
056
056
057
052
053
042
062
061
067
088
056
061
076
063
070
057
060
043
032
055
062
069
062
067
068
053
052
052
039
067
057
064
051
073
064
087
047
057
060
Austropurcelliascoparia
055
050
054
052
053
067
052
068
054
051
045
065
050
064
057
044
071
065
074
059
071
072
051
054
084
053
052
052
071
052
082
055
088
062
056
056
061
091
053
Austrop
urcelliawoo
dwardi
054
065
058
052
058
056
061
069
053
053
042
066
049
066
068
053
057
062
068
054
063
047
049
036
050
067
052
060
056
065
079
064
068
059
056
056
047
061
053
Chileogovea
jocasta
049
053
055
052
054
058
050
057
055
049
047
049
058
060
055
063
050
050
058
051
074
055
053
053
060
053
052
055
061
057
069
057
068
066
082
088
082
053
069
Chileog
ovea
Oedipus
070
053
054
052
053
064
070
074
057
062
053
052
043
064
054
051
058
072
078
053
071
012
049
043
054
052
052
052
061
052
063
061
070
062
092
091
083
051
048
Pettaluslampetid
es063
066
078
052
092
067
057
058
053
055
063
039
048
055
057
063
064
068
068
046
064
050
095
090
044
084
052
093
065
042
050
052
066
074
056
056
047
052
055
Pettaluscimicifo
rmis
100
110
140
066
095
089
082
076
066
058
074
033
039
021
057
058
058
056
046
041
056
085
125
064
044
116
059
074
070
039
047
055
071
068
056
056
047
050
061
Troglosiroaelleni
059
053
059
054
066
066
064
057
054
053
052
040
067
066
072
060
064
044
053
076
045
064
066
075
062
054
054
067
068
054
061
066
064
074
056
056
047
055
058
Troglosirolong
ifossa
051
045
055
052
073
064
048
070
049
060
057
042
057
053
069
043
069
053
064
079
041
059
055
061
066
052
052
081
070
049
049
052
055
074
056
056
047
056
053
Troglosironinq
ua059
045
057
053
086
068
049
074
049
066
048
044
074
054
064
052
063
048
059
076
044
054
063
072
068
052
054
092
069
062
076
048
070
074
056
056
047
057
072
Huitaca
ventralis
075
045
057
052
060
036
084
047
105
094
063
054
058
067
072
068
054
040
056
075
057
046
058
070
053
052
052
058
039
045
038
057
047
074
056
056
047
052
048
Neogo
veasp
067
056
058
065
070
031
070
075
081
099
042
055
067
047
063
068
042
031
045
074
050
038
061
073
061
052
063
069
033
035
035
055
044
074
056
056
047
050
045
Metagovea
philipi
045
045
054
052
053
041
045
057
065
055
056
056
066
051
053
054
056
048
062
078
057
057
053
050
031
052
052
052
036
042
056
048
073
074
056
056
047
053
068
Cypho
phthalmus
duricorius
050
045
055
052
053
058
041
050
039
060
067
044
044
042
027
050
072
058
065
078
039
065
057
071
065
052
052
052
062
059
067
044
072
074
056
056
047
056
065
Parasirocoiffaiti
044
045
055
054
055
058
056
047
050
052
079
050
041
060
056
080
051
064
061
086
042
064
060
062
058
052
056
057
068
053
052
081
063
068
056
056
055
053
064
Siro
acaroids
046
055
054
052
053
055
053
049
044
055
047
051
067
060
070
051
057
058
071
069
048
068
053
044
061
053
052
052
058
062
061
062
055
051
056
056
071
055
068
Siro
rubens
049
045
056
052
053
042
077
050
041
055
065
039
057
053
062
082
037
047
052
072
035
073
059
052
086
052
052
052
036
054
085
102
100
066
119
102
056
050
049
Suzukielus
sauteri
059
045
056
052
053
060
062
048
050
051
063
068
069
065
064
066
056
082
071
075
048
071
056
067
057
052
052
052
062
079
072
066
063
071
098
091
047
056
061
Stylocellusramblae
052
045
055
052
053
064
050
060
052
048
083
058
047
056
053
055
076
045
046
092
068
060
054
046
048
052
052
052
067
053
058
053
074
074
056
056
047
053
060
Stylocellustambusisi
074
045
057
052
053
066
049
059
046
067
058
052
050
041
029
059
076
047
056
082
066
060
057
070
056
052
052
052
061
047
055
041
065
074
056
056
047
052
062
Fan
gensiscavernarus
081
064
065
052
053
073
046
047
049
065
080
054
076
053
065
048
079
048
026
075
061
073
065
078
081
059
052
052
066
063
049
038
051
074
056
056
047
053
048
Fan
gensisspelaeus
073
045
057
052
053
078
093
047
050
061
093
065
077
056
064
076
053
068
030
076
054
069
056
069
066
052
052
052
076
068
055
086
061
074
056
056
047
057
054
374 Invertebrate Systematics B L de Bivort and G Giribet
4041
4243
4445
4647
4849
5051
5253
5455
5657
5859
6061
6263
6465
6667
6869
7071
7273
7475
7677
78
Parap
urcelliastaregaisp
nov
054
049
048
053
061
054
046
048
065
057
060
056
057
059
069
055
074
063
048
049
055
058
069
062
068
080
057
066
053
057
056
052
068
059
066
054
070
051
062
Parap
urcelliaam
atolaesp
nov
047
058
055
044
066
054
046
050
081
058
068
069
065
064
068
055
068
055
049
032
047
064
064
070
066
059
053
073
046
043
035
075
066
053
068
048
065
062
056
Parapurcelliaconvexa
spn
ov
036
030
068
095
053
054
055
050
064
057
053
052
056
065
070
055
069
066
055
062
053
069
052
075
070
059
063
067
049
055
069
062
077
034
062
057
071
052
062
Parapurcellianatalia
sp
nov
050
046
050
051
054
054
046
046
079
060
056
066
072
063
064
055
062
060
057
058
046
068
053
063
063
069
042
063
045
054
073
050
062
073
056
054
060
048
054
Parap
urcelliaminuta
spn
ov
046
046
054
041
030
054
047
049
075
060
059
050
058
059
066
055
062
062
057
040
062
066
072
061
062
076
044
064
052
056
058
076
095
069
053
058
062
057
054
Parap
urcelliaperegrinator
058
058
058
050
049
054
046
054
071
063
049
056
070
062
060
055
062
060
057
070
053
063
073
077
079
056
052
078
049
061
058
057
064
079
052
048
066
055
055
Parapurcelliasilvicola
052
054
062
050
044
054
050
048
064
058
063
059
055
062
069
055
064
061
052
051
062
067
072
070
066
019
045
064
059
059
048
061
074
066
065
073
070
064
057
Parapurcelliamonticola
067
073
086
060
055
054
053
051
066
057
050
062
071
063
070
055
060
065
058
053
044
064
061
039
072
075
047
080
048
061
056
047
066
061
066
047
061
053
056
Parap
urcelliarumpian
a040
035
072
081
058
054
055
045
075
057
048
067
074
063
065
055
082
059
056
072
052
066
051
064
073
068
049
076
073
066
056
058
070
035
054
090
078
093
058
Parapurcelliafissa
060
057
065
048
059
054
057
060
063
057
049
056
066
063
060
055
062
061
051
064
056
065
062
038
074
051
057
079
050
059
043
048
069
061
067
051
071
055
060
Purcelliatransvaalica
064
068
060
051
053
054
056
057
062
055
044
054
065
055
069
055
077
068
052
054
044
068
069
066
064
058
060
058
060
071
050
043
054
051
073
055
054
056
058
Purcelliaillustrans
111
080
101
074
037
054
082
067
066
052
047
046
060
066
057
055
056
056
055
047
052
066
063
057
066
048
055
077
052
068
051
043
059
048
053
047
072
054
059
Purcelliagriswoldisp
nov
067
078
063
048
049
054
054
049
065
057
045
050
066
061
065
055
053
061
059
046
050
063
074
056
070
071
070
066
058
070
035
050
051
069
049
058
077
052
056
Purcellialawrenceisp
nov
071
081
067
052
046
054
056
052
089
054
046
040
054
061
066
055
059
059
069
047
038
074
067
054
072
068
056
056
056
065
051
050
050
050
042
062
068
052
057
Speleosiro
argasiform
is057
056
054
045
060
054
047
044
055
066
041
058
078
061
045
055
058
061
057
044
076
049
090
079
055
054
085
058
077
089
050
052
051
070
043
041
050
048
060
Rakaiaisolate
066
075
061
060
062
054
061
079
032
073
059
069
071
063
059
055
066
060
066
044
061
052
039
053
069
056
038
058
049
050
049
077
071
058
077
062
065
064
059
Rakaialin
dsayi
050
060
058
056
068
054
057
054
070
057
067
065
060
067
060
055
055
056
068
059
073
059
060
044
078
074
049
053
060
059
050
064
057
085
059
055
064
066
058
Rakaiamag
naau
stralis
060
047
055
055
055
054
046
052
059
078
064
074
072
066
054
055
061
064
073
060
060
064
050
056
070
047
058
068
062
052
053
074
049
049
058
061
070
077
058
Rakaiamedia
060
082
041
052
062
054
061
052
065
078
064
082
075
061
058
055
060
062
075
060
047
059
043
059
063
071
060
064
049
037
076
056
070
050
061
063
070
051
060
Rakaiasolitaria
065
071
056
052
051
054
060
066
068
067
074
075
063
069
053
055
078
056
065
060
053
064
047
055
063
051
057
081
050
048
074
072
056
065
075
069
073
052
058
Rakaiasorensenidigitata
076
071
066
060
034
054
087
052
064
060
049
030
039
084
055
055
065
065
065
060
068
057
072
072
072
051
067
069
061
062
061
078
076
078
077
073
071
070
048
Aorakicrypta
071
069
053
060
054
054
082
048
068
067
068
063
051
069
055
055
059
073
071
054
064
047
075
078
071
056
080
040
068
049
067
081
042
064
068
041
047
057
048
Aorakidenticula
075
068
054
061
068
054
085
055
052
057
068
055
048
077
054
055
070
060
056
065
047
073
071
049
074
066
053
055
057
035
085
061
070
067
058
040
059
054
060
Aorakiinerma
069
070
071
078
065
054
069
046
068
060
064
063
052
072
050
055
081
064
054
050
058
060
062
072
042
055
057
073
070
068
054
052
047
043
079
046
057
055
061
Aorakilong
itarsa
069
066
057
062
073
054
067
078
064
059
075
063
056
071
056
055
065
073
057
069
073
043
074
071
064
050
071
054
064
049
071
078
050
052
062
056
063
057
062
Aorakitumidata
066
062
067
047
063
054
073
062
066
055
074
087
071
070
065
055
051
065
072
062
095
023
074
082
079
081
077
063
082
060
064
105
068
085
055
059
071
069
069
Karripurcellia
harveyi
050
061
072
074
086
054
084
076
057
054
047
047
064
062
057
055
056
054
061
077
075
053
060
073
053
047
066
059
064
070
054
051
055
061
045
052
054
050
060
Karripurcellia
peckorum
046
050
066
077
087
054
067
056
050
067
045
047
062
064
056
055
059
059
074
071
069
056
061
059
063
045
055
049
082
067
054
055
042
054
081
071
051
090
055
Karripurcellia
sierwaldae
050
052
064
067
068
054
062
061
060
050
060
044
044
064
065
055
052
060
051
060
060
058
057
064
061
060
063
070
079
074
051
040
045
058
056
081
057
097
048
Austropurcelliascoparia
079
078
069
066
053
054
074
064
051
052
080
056
037
089
038
055
056
057
080
050
057
076
067
062
060
048
053
056
052
062
064
060
061
068
062
066
062
052
063
Austrop
urcelliawoo
dwardi
072
067
074
061
054
054
075
079
049
062
075
067
047
074
036
055
058
058
053
054
051
086
061
052
056
059
038
054
054
065
060
046
058
049
063
080
067
054
048
Chileogovea
jocasta
067
064
065
054
054
054
061
052
054
055
069
066
052
069
045
055
049
058
057
057
063
066
042
053
059
068
068
059
056
074
054
056
057
060
043
057
046
050
048
Chileog
ovea
oedipu
s064
068
070
070
048
054
070
076
081
056
070
056
046
069
056
055
054
059
054
085
072
060
047
044
076
057
066
055
072
074
048
064
054
076
046
052
041
060
048
Pettaluslampetid
es063
055
059
058
058
054
057
051
063
063
053
077
084
052
075
055
058
051
057
058
054
052
051
057
060
063
062
052
073
067
054
051
039
075
083
040
034
053
061
Pettaluscimicifo
rmis
063
061
050
062
048
062
060
058
044
065
044
054
079
048
070
055
042
048
069
049
060
051
057
051
058
062
082
054
051
061
077
059
060
053
072
046
062
051
057
Troglosiroaelleni
059
056
029
089
066
085
058
054
066
046
078
071
050
062
045
090
056
060
068
076
055
055
066
058
045
064
076
054
064
065
059
049
055
055
046
049
037
052
060
Troglosirolongifo
ssa
059
055
058
070
077
085
077
053
061
045
069
071
058
048
054
093
038
065
070
085
059
056
052
082
044
051
071
049
056
045
075
053
050
053
038
063
073
064
048
Troglosironinq
ua066
051
072
073
079
083
077
061
060
041
068
067
053
045
064
092
062
059
058
067
059
053
070
075
052
061
082
049
075
076
046
055
048
051
036
053
045
068
048
Huitaca
ventralis
045
045
040
050
059
085
046
058
056
041
049
070
083
032
075
081
031
044
049
051
062
061
070
030
046
062
071
090
108
096
085
062
043
073
043
063
033
084
048
Neogo
veasp
042
067
051
086
073
087
071
066
031
046
039
066
087
037
059
055
066
130
104
061
066
071
062
039
042
072
075
076
074
077
079
055
050
036
084
064
038
095
061
Metag
ovea
philipi
057
068
061
073
063
085
062
044
040
041
066
069
059
039
057
055
061
044
080
064
056
085
058
045
035
059
041
075
059
063
053
049
046
047
065
054
034
059
048
Cypho
phthalmus
duricorius
060
046
053
056
061
086
050
058
065
056
053
064
073
054
051
087
060
064
057
082
060
061
060
065
048
047
064
036
055
052
070
059
066
065
071
077
072
064
078
Parasirocoiffaiti
056
064
071
050
066
081
046
074
063
060
089
070
051
051
055
078
055
065
053
062
063
060
032
073
042
079
053
052
044
048
078
068
088
061
060
077
070
055
048
Siro
acaroides
052
040
038
053
081
085
049
077
058
060
057
079
073
055
059
096
032
061
049
068
045
071
040
057
055
059
058
043
065
064
063
065
066
050
059
067
056
059
048
Siro
rubens
063
050
064
053
063
085
050
069
056
049
076
069
054
060
051
087
073
067
078
089
056
064
034
060
046
084
064
038
056
054
068
064
076
049
045
063
068
058
073
Suzukielus
sauteri
064
047
063
048
073
054
047
101
069
053
063
050
048
052
075
055
060
062
067
077
059
058
042
069
068
069
052
050
049
048
075
061
066
056
069
065
074
050
068
Stylocellusramblae
060
072
062
053
058
054
062
073
041
098
074
042
040
035
071
055
025
044
038
053
105
053
072
051
030
061
082
052
056
052
067
075
059
072
063
066
048
047
099
Stylocellustambusisi
056
063
061
052
060
054
056
069
047
096
073
050
041
040
067
055
063
044
038
042
064
060
074
050
036
048
063
044
060
049
067
060
058
078
057
085
046
062
087
Fan
gensiscavernarus
047
044
040
056
075
054
046
070
036
085
050
042
058
041
084
055
077
044
038
066
076
035
052
064
049
055
070
046
051
044
056
067
062
071
050
058
060
057
096
Fan
gensisspelaeus
049
061
044
055
079
054
046
070
047
086
046
035
058
042
105
055
087
044
038
065
077
029
068
065
060
084
050
047
051
045
078
070
083
057
054
071
065
056
096
Systematic revision of South African pettalids Invertebrate Systematics 375
combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)
Methods and abbreviations
Specimens used in this study were provided by the followinginstitutions
AMNH American Museum of Natural History New York(USA)
BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard
University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand
Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo
Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg
(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg
Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)
Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg
(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen
Copenhagen (Denmark)
Specimen handling and illustration
In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was
photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy
Character coding
Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3
Phylogenetic analyses
The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)
The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the
376 Invertebrate Systematics B L de Bivort and G Giribet
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
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A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
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H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
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lyfork
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Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
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a r
um
pia
na
Ra
kaia
iso
lata
Pu
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gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
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lo
ng
ita
rsa
Pu
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law
ren
cei s
p n
ov
Au
stro
pu
rce
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sco
pa
ria
Ka
rrip
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a p
eck
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mK
arr
ipu
rce
llia
ha
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yi
Pa
rap
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a m
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ta s
p n
ov
Ra
kaia
so
ren
sen
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itata
Ka
rrip
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a s
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ald
ae
Pe
tta
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cim
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rmis
Pa
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urc
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a p
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tor
Ra
kaia
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a a
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Pa
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a s
tare
ga
i sp
nov
Sir
o r
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Pa
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Tro
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Pa
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Tro
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i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
4041
4243
4445
4647
4849
5051
5253
5455
5657
5859
6061
6263
6465
6667
6869
7071
7273
7475
7677
78
Parap
urcelliastaregaisp
nov
054
049
048
053
061
054
046
048
065
057
060
056
057
059
069
055
074
063
048
049
055
058
069
062
068
080
057
066
053
057
056
052
068
059
066
054
070
051
062
Parap
urcelliaam
atolaesp
nov
047
058
055
044
066
054
046
050
081
058
068
069
065
064
068
055
068
055
049
032
047
064
064
070
066
059
053
073
046
043
035
075
066
053
068
048
065
062
056
Parapurcelliaconvexa
spn
ov
036
030
068
095
053
054
055
050
064
057
053
052
056
065
070
055
069
066
055
062
053
069
052
075
070
059
063
067
049
055
069
062
077
034
062
057
071
052
062
Parapurcellianatalia
sp
nov
050
046
050
051
054
054
046
046
079
060
056
066
072
063
064
055
062
060
057
058
046
068
053
063
063
069
042
063
045
054
073
050
062
073
056
054
060
048
054
Parap
urcelliaminuta
spn
ov
046
046
054
041
030
054
047
049
075
060
059
050
058
059
066
055
062
062
057
040
062
066
072
061
062
076
044
064
052
056
058
076
095
069
053
058
062
057
054
Parap
urcelliaperegrinator
058
058
058
050
049
054
046
054
071
063
049
056
070
062
060
055
062
060
057
070
053
063
073
077
079
056
052
078
049
061
058
057
064
079
052
048
066
055
055
Parapurcelliasilvicola
052
054
062
050
044
054
050
048
064
058
063
059
055
062
069
055
064
061
052
051
062
067
072
070
066
019
045
064
059
059
048
061
074
066
065
073
070
064
057
Parapurcelliamonticola
067
073
086
060
055
054
053
051
066
057
050
062
071
063
070
055
060
065
058
053
044
064
061
039
072
075
047
080
048
061
056
047
066
061
066
047
061
053
056
Parap
urcelliarumpian
a040
035
072
081
058
054
055
045
075
057
048
067
074
063
065
055
082
059
056
072
052
066
051
064
073
068
049
076
073
066
056
058
070
035
054
090
078
093
058
Parapurcelliafissa
060
057
065
048
059
054
057
060
063
057
049
056
066
063
060
055
062
061
051
064
056
065
062
038
074
051
057
079
050
059
043
048
069
061
067
051
071
055
060
Purcelliatransvaalica
064
068
060
051
053
054
056
057
062
055
044
054
065
055
069
055
077
068
052
054
044
068
069
066
064
058
060
058
060
071
050
043
054
051
073
055
054
056
058
Purcelliaillustrans
111
080
101
074
037
054
082
067
066
052
047
046
060
066
057
055
056
056
055
047
052
066
063
057
066
048
055
077
052
068
051
043
059
048
053
047
072
054
059
Purcelliagriswoldisp
nov
067
078
063
048
049
054
054
049
065
057
045
050
066
061
065
055
053
061
059
046
050
063
074
056
070
071
070
066
058
070
035
050
051
069
049
058
077
052
056
Purcellialawrenceisp
nov
071
081
067
052
046
054
056
052
089
054
046
040
054
061
066
055
059
059
069
047
038
074
067
054
072
068
056
056
056
065
051
050
050
050
042
062
068
052
057
Speleosiro
argasiform
is057
056
054
045
060
054
047
044
055
066
041
058
078
061
045
055
058
061
057
044
076
049
090
079
055
054
085
058
077
089
050
052
051
070
043
041
050
048
060
Rakaiaisolate
066
075
061
060
062
054
061
079
032
073
059
069
071
063
059
055
066
060
066
044
061
052
039
053
069
056
038
058
049
050
049
077
071
058
077
062
065
064
059
Rakaialin
dsayi
050
060
058
056
068
054
057
054
070
057
067
065
060
067
060
055
055
056
068
059
073
059
060
044
078
074
049
053
060
059
050
064
057
085
059
055
064
066
058
Rakaiamag
naau
stralis
060
047
055
055
055
054
046
052
059
078
064
074
072
066
054
055
061
064
073
060
060
064
050
056
070
047
058
068
062
052
053
074
049
049
058
061
070
077
058
Rakaiamedia
060
082
041
052
062
054
061
052
065
078
064
082
075
061
058
055
060
062
075
060
047
059
043
059
063
071
060
064
049
037
076
056
070
050
061
063
070
051
060
Rakaiasolitaria
065
071
056
052
051
054
060
066
068
067
074
075
063
069
053
055
078
056
065
060
053
064
047
055
063
051
057
081
050
048
074
072
056
065
075
069
073
052
058
Rakaiasorensenidigitata
076
071
066
060
034
054
087
052
064
060
049
030
039
084
055
055
065
065
065
060
068
057
072
072
072
051
067
069
061
062
061
078
076
078
077
073
071
070
048
Aorakicrypta
071
069
053
060
054
054
082
048
068
067
068
063
051
069
055
055
059
073
071
054
064
047
075
078
071
056
080
040
068
049
067
081
042
064
068
041
047
057
048
Aorakidenticula
075
068
054
061
068
054
085
055
052
057
068
055
048
077
054
055
070
060
056
065
047
073
071
049
074
066
053
055
057
035
085
061
070
067
058
040
059
054
060
Aorakiinerma
069
070
071
078
065
054
069
046
068
060
064
063
052
072
050
055
081
064
054
050
058
060
062
072
042
055
057
073
070
068
054
052
047
043
079
046
057
055
061
Aorakilong
itarsa
069
066
057
062
073
054
067
078
064
059
075
063
056
071
056
055
065
073
057
069
073
043
074
071
064
050
071
054
064
049
071
078
050
052
062
056
063
057
062
Aorakitumidata
066
062
067
047
063
054
073
062
066
055
074
087
071
070
065
055
051
065
072
062
095
023
074
082
079
081
077
063
082
060
064
105
068
085
055
059
071
069
069
Karripurcellia
harveyi
050
061
072
074
086
054
084
076
057
054
047
047
064
062
057
055
056
054
061
077
075
053
060
073
053
047
066
059
064
070
054
051
055
061
045
052
054
050
060
Karripurcellia
peckorum
046
050
066
077
087
054
067
056
050
067
045
047
062
064
056
055
059
059
074
071
069
056
061
059
063
045
055
049
082
067
054
055
042
054
081
071
051
090
055
Karripurcellia
sierwaldae
050
052
064
067
068
054
062
061
060
050
060
044
044
064
065
055
052
060
051
060
060
058
057
064
061
060
063
070
079
074
051
040
045
058
056
081
057
097
048
Austropurcelliascoparia
079
078
069
066
053
054
074
064
051
052
080
056
037
089
038
055
056
057
080
050
057
076
067
062
060
048
053
056
052
062
064
060
061
068
062
066
062
052
063
Austrop
urcelliawoo
dwardi
072
067
074
061
054
054
075
079
049
062
075
067
047
074
036
055
058
058
053
054
051
086
061
052
056
059
038
054
054
065
060
046
058
049
063
080
067
054
048
Chileogovea
jocasta
067
064
065
054
054
054
061
052
054
055
069
066
052
069
045
055
049
058
057
057
063
066
042
053
059
068
068
059
056
074
054
056
057
060
043
057
046
050
048
Chileog
ovea
oedipu
s064
068
070
070
048
054
070
076
081
056
070
056
046
069
056
055
054
059
054
085
072
060
047
044
076
057
066
055
072
074
048
064
054
076
046
052
041
060
048
Pettaluslampetid
es063
055
059
058
058
054
057
051
063
063
053
077
084
052
075
055
058
051
057
058
054
052
051
057
060
063
062
052
073
067
054
051
039
075
083
040
034
053
061
Pettaluscimicifo
rmis
063
061
050
062
048
062
060
058
044
065
044
054
079
048
070
055
042
048
069
049
060
051
057
051
058
062
082
054
051
061
077
059
060
053
072
046
062
051
057
Troglosiroaelleni
059
056
029
089
066
085
058
054
066
046
078
071
050
062
045
090
056
060
068
076
055
055
066
058
045
064
076
054
064
065
059
049
055
055
046
049
037
052
060
Troglosirolongifo
ssa
059
055
058
070
077
085
077
053
061
045
069
071
058
048
054
093
038
065
070
085
059
056
052
082
044
051
071
049
056
045
075
053
050
053
038
063
073
064
048
Troglosironinq
ua066
051
072
073
079
083
077
061
060
041
068
067
053
045
064
092
062
059
058
067
059
053
070
075
052
061
082
049
075
076
046
055
048
051
036
053
045
068
048
Huitaca
ventralis
045
045
040
050
059
085
046
058
056
041
049
070
083
032
075
081
031
044
049
051
062
061
070
030
046
062
071
090
108
096
085
062
043
073
043
063
033
084
048
Neogo
veasp
042
067
051
086
073
087
071
066
031
046
039
066
087
037
059
055
066
130
104
061
066
071
062
039
042
072
075
076
074
077
079
055
050
036
084
064
038
095
061
Metag
ovea
philipi
057
068
061
073
063
085
062
044
040
041
066
069
059
039
057
055
061
044
080
064
056
085
058
045
035
059
041
075
059
063
053
049
046
047
065
054
034
059
048
Cypho
phthalmus
duricorius
060
046
053
056
061
086
050
058
065
056
053
064
073
054
051
087
060
064
057
082
060
061
060
065
048
047
064
036
055
052
070
059
066
065
071
077
072
064
078
Parasirocoiffaiti
056
064
071
050
066
081
046
074
063
060
089
070
051
051
055
078
055
065
053
062
063
060
032
073
042
079
053
052
044
048
078
068
088
061
060
077
070
055
048
Siro
acaroides
052
040
038
053
081
085
049
077
058
060
057
079
073
055
059
096
032
061
049
068
045
071
040
057
055
059
058
043
065
064
063
065
066
050
059
067
056
059
048
Siro
rubens
063
050
064
053
063
085
050
069
056
049
076
069
054
060
051
087
073
067
078
089
056
064
034
060
046
084
064
038
056
054
068
064
076
049
045
063
068
058
073
Suzukielus
sauteri
064
047
063
048
073
054
047
101
069
053
063
050
048
052
075
055
060
062
067
077
059
058
042
069
068
069
052
050
049
048
075
061
066
056
069
065
074
050
068
Stylocellusramblae
060
072
062
053
058
054
062
073
041
098
074
042
040
035
071
055
025
044
038
053
105
053
072
051
030
061
082
052
056
052
067
075
059
072
063
066
048
047
099
Stylocellustambusisi
056
063
061
052
060
054
056
069
047
096
073
050
041
040
067
055
063
044
038
042
064
060
074
050
036
048
063
044
060
049
067
060
058
078
057
085
046
062
087
Fan
gensiscavernarus
047
044
040
056
075
054
046
070
036
085
050
042
058
041
084
055
077
044
038
066
076
035
052
064
049
055
070
046
051
044
056
067
062
071
050
058
060
057
096
Fan
gensisspelaeus
049
061
044
055
079
054
046
070
047
086
046
035
058
042
105
055
087
044
038
065
077
029
068
065
060
084
050
047
051
045
078
070
083
057
054
071
065
056
096
Systematic revision of South African pettalids Invertebrate Systematics 375
combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)
Methods and abbreviations
Specimens used in this study were provided by the followinginstitutions
AMNH American Museum of Natural History New York(USA)
BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard
University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand
Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo
Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg
(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg
Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)
Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg
(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen
Copenhagen (Denmark)
Specimen handling and illustration
In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was
photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy
Character coding
Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3
Phylogenetic analyses
The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)
The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the
376 Invertebrate Systematics B L de Bivort and G Giribet
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
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F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)
Methods and abbreviations
Specimens used in this study were provided by the followinginstitutions
AMNH American Museum of Natural History New York(USA)
BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard
University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand
Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo
Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg
(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg
Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)
Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg
(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen
Copenhagen (Denmark)
Specimen handling and illustration
In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was
photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy
Character coding
Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3
Phylogenetic analyses
The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)
The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the
376 Invertebrate Systematics B L de Bivort and G Giribet
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
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A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
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Fig11
(A)S
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Systematic revision of South African pettalids Invertebrate Systematics 397
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
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ingfrom
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dinwhich
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pwith
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ithinthePettalid
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ndstoalltreesun
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ata
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umberson
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analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data
Taxonomy
Order OPILIONES Sundevall
Suborder CYPHOPHTHALMI Simon
Family PETTALIDAE Simon
Genus Speleosiro Lawrence 1931
Diagnosis
Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII
Type species
Speleosiro argasiformis Lawrence 1931
Species included
Speleosiro argasiformis Lawrence 1931
Distribution
SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province
Remarks
This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement
Genus Purcellia Hansen amp Soslashrensen 1904
Diagnosis
Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)
Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)
Type species
Purcellia illustrans Hansen amp Soslashrensen 1904
Species included
Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov
Distribution
South Africa Limpopo Eastern Cape and Western CapeProvinces
Remarks
This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa
Purcellia griswoldi sp nov
(Figs 2 3 Table 3)
Material examined
Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985
Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981
Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981
Systematic revision of South African pettalids Invertebrate Systematics 377
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
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oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
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lyforkrangingfrom
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Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
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onca
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(k)
poly
phyl
etic
para
phyl
etic
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letic
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45
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21
11
12
14
18
Pet
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ae S
tylo
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Neo
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Tro
glos
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(T
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Aus
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ea P
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lus
(Pur
celli
a gr
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oldi
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nov
ampP
urce
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renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
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celli
a)
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celli
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leos
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(Pur
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Spe
leos
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Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
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me
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Pa
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um
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Ra
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Pu
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gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
A B
E
F G H
I
F G H
I
N O P
Q
N O P
R S TR S TQ
J K L
M
J K L
M
DC
378 Invertebrate Systematics B L de Bivort and G Giribet
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Diagnosis
With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view
Description
Male
Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV
endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites
Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)
Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral
A B
Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm
Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm
Systematic revision of South African pettalids Invertebrate Systematics 379
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
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Systematic revision of South African pettalids Invertebrate Systematics 397
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Fig12
(A)Navajorugs
summarisingresults
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onyusingcombineddiscreteandcontinuous
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hesearch
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ithinthePettalid
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ata
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umberson
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=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long
Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all
trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length
Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged
Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture
Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum
Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092
Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756
Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696
Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722
Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344
Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637
Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779
380 Invertebrate Systematics B L de Bivort and G Giribet
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
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Fig11
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Systematic revision of South African pettalids Invertebrate Systematics 397
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
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ingfrom
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dinwhich
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pwith
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ithinthePettalid
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ndstoalltreesun
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ata
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umberson
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weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor
Female
Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess
Remarks
This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality
Distribution
Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east
Habitat
Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest
Etymology
This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids
Purcellia lawrencei sp nov
(Fig 4 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the
following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939
Paratype 1 (NMSA) same collecting data as holotype
Diagnosis
Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view
Description
Male
Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area
Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites
Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites
Systematic revision of South African pettalids Invertebrate Systematics 381
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
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Fig11
(A)S
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tained
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yanalysisofthediscretedatasetund
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Systematic revision of South African pettalids Invertebrate Systematics 397
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99 51 69
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Fig12
(A)Navajorugs
summarisingresults
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onyusingcombineddiscreteandcontinuous
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hesearch
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acrossequalw
eights(EW)and
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ingfrom
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trictconsensus
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foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
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derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
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11
11
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14
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ofalltreesinwhich
thegenusParapurcelliaisfoun
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basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
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and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
AP
sc
opTIX
S9 S8S7
S6
TVIII
S5
C
I J K L
A B
D E
CIV
S1
G
CI
CII
CIII
EP
ECIII
ECIVCG
ECI
ECII
F G HF G H
382 Invertebrate Systematics B L de Bivort and G Giribet
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
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M N O PN O P
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F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)
Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles
Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long
Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length
Spermatopositor not studied due to the single male specimenavailable (the holotype)
Female
Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline
Remarks
This species is likely sympatric with Purcellia griswoldi
Distribution
Only known from the type locality near Knysna Western CapeProvince South Africa
Habitat
No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest
Etymology
The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods
Genus Parapurcellia Rosas Costa 1950
Diagnosis
Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)
Type species
Parapurcellia fissa (Lawrence 1939)
Species included
Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov
Distribution
South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces
Remarks
This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa
Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm
Systematic revision of South African pettalids Invertebrate Systematics 383
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
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valuesrang
ingfrom
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Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
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(k)
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etic
para
phyl
etic
mon
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letic
approx tree length contributionratio (continuousdiscrete)
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41
21
11
12
14
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Pet
talid
ae S
tylo
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Neo
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Tro
glos
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dae
(T
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Neo
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idae
)K
arrip
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Aus
trop
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C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
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sp
nov
ampP
urce
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law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
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(Par
apur
celli
a)
(Pur
celli
a +
Par
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Spe
leos
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(Pur
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a +
Par
apur
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a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
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de
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cula
Su
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s sa
ute
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Ra
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me
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Pa
rap
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swo
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a t
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Ka
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i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
C
A B
D E
F G H IF G H I
N O P QN O P Q
J K L MJ K L M
384 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
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Fig11
(A)S
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tained
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yanalysisofthediscretedatasetund
erim
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Systematic revision of South African pettalids Invertebrate Systematics 397
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
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ingfrom
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trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
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ophyletic)Thiscorrespon
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derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
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11
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and18(C)S
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ofalltreesinwhich
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ithinthePettalid
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ndstoalltreesun
derk
=1andk=2andk=3ford
ata
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sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Parapurcellia amatola sp nov
(Fig 5 Table 3)
Material examined
Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965
Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting
data as holotype
Diagnosis
Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline
Description
Male
Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)
Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit
Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long
Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length
Spermatopositor not studied
Female
Unknown
Distribution
Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province
Habitat
No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest
Etymology
Amatola a noun in apposition after its collection locality theAmatola forest
Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm
Systematic revision of South African pettalids Invertebrate Systematics 385
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
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oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
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lyforkrangingfrom
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Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
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onca
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(k)
poly
phyl
etic
para
phyl
etic
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letic
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45
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21
11
12
14
18
Pet
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ae S
tylo
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Neo
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Tro
glos
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(T
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Aus
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ea P
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lus
(Pur
celli
a gr
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oldi
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nov
ampP
urce
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renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
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celli
a)
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celli
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leos
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(Pur
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Spe
leos
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Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
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me
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Pa
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um
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Ra
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Pu
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gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
C
A B
D E
H
I
F G H
I J K L
M
J K L
M N O PN O P
386 Invertebrate Systematics B L de Bivort and G Giribet
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
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idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
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Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
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Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
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pta
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cula
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ute
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ipu
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ren
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itata
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a s
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a p
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gn
a a
ust
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Pa
rap
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elli
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tare
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i sp
nov
Sir
o r
ub
en
s
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ro c
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i
Tro
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ata
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Pa
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elli
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Ne
og
ove
a s
p
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elli
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issa
Tro
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ifoss
a
Pa
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elli
a m
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la
Pe
tta
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lam
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Ao
raki
tu
mid
ata
Me
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a p
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pi
Au
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llia
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od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
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qu
a
Pa
rap
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elli
a s
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Ao
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erm
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Cyp
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uri
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us
Sp
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Ch
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us
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en
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Fa
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Fa
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Fa
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rum
Pu
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Ao
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tu
mid
ata
Pe
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lus
lam
pe
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es
Me
tag
ove
a p
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pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
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lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
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llia
sco
pa
ria
Ka
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elli
a p
eck
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m
Pa
rap
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elli
a a
ma
tola
sp
nov
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llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
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elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
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ov
Pa
rap
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elli
a s
tare
ga
i sp
nov
Pe
tta
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icifo
rmis
Pa
rap
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elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Parapurcellia convexa sp nov
(Fig 6 Table 3)
Material examined
Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985
Paratypes 3 lt 1 (NMSA) same collecting data as holotype
Diagnosis
Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated
Description
Male
Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)
Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules
Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)
Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long
Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-
Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm
Systematic revision of South African pettalids Invertebrate Systematics 387
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
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Neo
gove
idae
Tro
glos
ironi
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(T
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Neo
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)K
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trop
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ogov
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(Pur
celli
a gr
isw
oldi
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nov
ampP
urce
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renc
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p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
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aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
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a +
Par
apur
celli
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Spe
leos
iro)
(Pur
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Par
apur
celli
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leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
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de
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cula
Su
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elu
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ute
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Ra
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ren
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itata
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ub
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Me
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igit
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o a
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Tro
glo
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a Ka
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a h
arv
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99 51 69
75 92 95 8581
80 69
94
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74 83
86 86 5665
6379
8877 100
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81 87
68
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
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mon
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dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
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11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
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ithinthePettalid
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paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
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sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa
Habitat
Specimens were collected in Podocarpus litter
Etymology
From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus
Parapurcellia minuta sp nov
(Fig 7 Table 3)
Material examined
Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940
Paratypes 1 lt 2 (NMSA) same collecting data as holotype
Diagnosis
Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)
Description
Male
Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)
Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)
Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long
Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented
Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm
388 Invertebrate Systematics B L de Bivort and G Giribet
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
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trict
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underparsim
onyanalysisof
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pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
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uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
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elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
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urc
elli
a f
issa
Tro
glo
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a
Pa
rap
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elli
a m
on
tico
la
Pe
tta
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lam
pe
tid
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Ao
raki
tu
mid
ata
Me
tag
ove
a p
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pi
Au
stro
pu
rce
llia
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wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
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Ao
raki
in
erm
a
Pu
rce
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stra
ns
Cyp
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ph
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s d
uri
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us
Sp
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Ch
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go
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us
Ra
kaia
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lita
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Hu
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ca v
en
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sta
Fa
ng
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rum
Sty
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bla
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Fa
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Fa
ng
en
sis
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Pu
rce
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stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
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es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
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sco
pa
ria
Ka
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a p
eck
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m
Pa
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ma
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sp
nov
Pu
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Ch
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us
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Ka
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ae
Pu
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aa
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Pa
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elli
a m
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ov
Pa
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Pe
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a f
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Fa
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en
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Tro
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Pa
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a s
ilvic
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Sty
loce
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tam
bu
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Pa
rap
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elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
C
II J K L
M N O PN O P
J K L
M
A B
D E
F G H
Systematic revision of South African pettalids Invertebrate Systematics 389
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
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lyfork
rangingfrom
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trict
consensusof
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underparsim
onyanalysisof
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pliedweightsforkvalues
rang
ingfrom
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)Asin
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uton
lyforkrangingfrom
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Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
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phyl
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phyl
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approx tree length contributionratio (continuousdiscrete)
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41
21
11
12
14
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Pet
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Neo
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glos
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talid
ae +
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aki +
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(Kar
ripur
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(Par
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leos
iro)
(Pur
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Aus
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urce
llia)
AB
C
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99 51 69
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
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continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
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dinwhich
thegenus
Parap
urcelliaisfoun
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pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length
Spermatopositor not studied
Female
Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline
Distribution
Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa
Habitat
No further information is available beyond the collection localitywhich has a mild subtropical climate
Etymology
From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length
Parapurcellia natalia sp nov
(Fig 8 Table 3)
Material examined
Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940
Diagnosis
Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin
Description
Male
Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without
lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)
Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)
Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long
Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing
390 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
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tained
underparsimon
yanalysisofthediscretedatasetund
erim
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Systematic revision of South African pettalids Invertebrate Systematics 397
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99 51 69
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
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mon
ophyletic)Thiscorrespon
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derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
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11
11
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and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
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ithinthePettalid
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paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
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weightin
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sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
C
F G H I
A B
D E
CI
CII
CIII
CIV
S1
G
ECII
ECIII
ECIVCG
EP
APsc
opTIX
S9
S8
S7
S6
TVIII
ECI
Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 391
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
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idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
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Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
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Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
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pta
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cula
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ute
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ipu
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ren
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itata
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a s
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a p
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gn
a a
ust
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Pa
rap
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elli
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tare
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i sp
nov
Sir
o r
ub
en
s
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ro c
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i
Tro
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ata
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Pa
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elli
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Ne
og
ove
a s
p
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elli
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issa
Tro
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ifoss
a
Pa
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elli
a m
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la
Pe
tta
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lam
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Ao
raki
tu
mid
ata
Me
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a p
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pi
Au
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llia
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od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
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qu
a
Pa
rap
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elli
a s
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Ao
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erm
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Cyp
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uri
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us
Sp
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Ch
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us
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en
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Fa
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Fa
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Fa
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rum
Pu
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Ao
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tu
mid
ata
Pe
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lus
lam
pe
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es
Me
tag
ove
a p
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pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
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lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
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llia
sco
pa
ria
Ka
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elli
a p
eck
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m
Pa
rap
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elli
a a
ma
tola
sp
nov
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llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
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elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
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ov
Pa
rap
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elli
a s
tare
ga
i sp
nov
Pe
tta
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icifo
rmis
Pa
rap
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elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
Distribution
Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected
Parapurcellia staregai sp nov
(Fig 9 Table 3)
Material examined
Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964
Diagnosis
Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed
Description
Male
Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma
Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other
Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)
Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)
Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each
Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long
Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length
Spermatopositor not studied as only the holotype wasavailable
Female
Unknown
392 Invertebrate Systematics B L de Bivort and G Giribet
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
C
F G H I
A B
D E
APsc
op
TIXS9
S8
S7
S6
TVIII
CI
CII
CIII
CIV
S1
G
ECIII
ECIVCG
EP
ECII
ECI
Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm
Systematic revision of South African pettalids Invertebrate Systematics 393
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
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Fig11
(A)S
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tained
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yanalysisofthediscretedatasetund
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Systematic revision of South African pettalids Invertebrate Systematics 397
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Fig12
(A)Navajorugs
summarisingresults
forcladesof
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underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
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ingfrom
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trictconsensus
ofalltrees
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dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
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derE
Wk=4k=5andk=6andk=3fordatapartition
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11
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and18(C)S
trictcon
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ithinthePettalid
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ndstoalltreesun
derk
=1andk=2andk=3ford
ata
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sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Distribution
Only known from the type locality in or near Trafalgar EasternCape Province South Africa
Habitat
No further information is available beyond the collection localitywhich has a subtropical moist savannah climate
Etymology
Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper
Results and discussion
The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove
In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae
Equally weighted analysis of each data partition
We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological
studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)
Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro
Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae
Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus
394 Invertebrate Systematics B L de Bivort and G Giribet
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree
The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus
The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola
Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B
Systematic revision of South African pettalids Invertebrate Systematics 395
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)
Implied weighting analysis of each data partition
Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only
The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia
For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia
Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied
weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30
Analysis of combined data under equaland implied weighting
The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)
Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly
Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the
396 Invertebrate Systematics B L de Bivort and G Giribet
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
AB
CD
Fig11
(A)S
trictcon
sensus
oftreesob
tained
underparsimon
yanalysisofthediscretedatasetund
erim
pliedweightsfork
valuesrang
ingfrom
1to6(B)A
sinAbuton
lyfork
rangingfrom
3to6(C)S
trict
consensusof
treesobtained
underparsim
onyanalysisof
thecontinuous
datasetunderim
pliedweightsforkvalues
rang
ingfrom
1to
6(D
)Asin
Cb
uton
lyforkrangingfrom
4to
6
Systematic revision of South African pettalids Invertebrate Systematics 397
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
EW
12
3
IW c
onca
vity
(k)
poly
phyl
etic
para
phyl
etic
mon
ophy
letic
approx tree length contributionratio (continuousdiscrete)
45
6
81
41
21
11
12
14
18
Pet
talid
ae S
tylo
celli
dae
Neo
gove
idae
Tro
glos
ironi
dae
(T
rogl
osiro
nida
e amp
Neo
gove
idae
)K
arrip
urce
llia
Aus
trop
urce
llia
C
hile
ogov
ea P
etta
lus
(Pur
celli
a gr
isw
oldi
sp
nov
ampP
urce
llia
law
renc
ei s
p n
ov)
(Pet
talid
ae +
Suz
ukie
lus)
(Aor
aki +
Rak
aia)
(Pet
talid
ae +
Siro
nida
e)
(Kar
ripur
celli
a +
Pet
talu
s)
(Par
apur
celli
a)
(Pur
celli
a +
Par
apur
celli
a+
Spe
leos
iro)
(Pur
celli
a +
Par
apur
celli
a +
Spe
leos
iro +
Aus
trop
urce
llia)
AB
C
Ao
raki
cry
pta
Ao
raki
de
nti
cula
Su
zuki
elu
s sa
ute
ri
Ra
kaia
me
dia
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
iso
lata
Pu
rce
llia
gri
swo
ldi
sp
no
vP
urc
elli
a t
ran
sva
alic
a
Ao
raki
lo
ng
ita
rsa
Pu
rce
llia
law
ren
cei s
p n
ov
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
mK
arr
ipu
rce
llia
ha
rve
yi
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Ra
kaia
so
ren
sen
i dig
itata
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a p
ere
gri
na
tor
Ra
kaia
ma
gn
a a
ust
ralis
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Sir
o r
ub
en
s
Pa
rasi
ro c
oif
fait
i
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Ne
og
ove
a s
p
Pa
rap
urc
elli
a f
issa
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a m
on
tico
la
Pe
tta
lus
lam
pe
tid
es
Ao
raki
tu
mid
ata
Me
tag
ove
a p
hili
pi
Au
stro
pu
rce
llia
wo
od
wa
rdi
Ra
kaia
lin
dsa
yi
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a
Pa
rap
urc
elli
a s
ilvic
ola
Ao
raki
in
erm
a
Pu
rce
llia
illu
stra
ns
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Sp
ele
osi
ro a
rga
sifo
rmis
Ch
ileo
go
vea
oe
dip
us
Ra
kaia
so
lita
ria
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Fa
ng
en
sis
cave
rna
rum
Sty
loce
llus
ram
bla
e
Fa
ng
en
sis
spe
lae
us
Sty
loce
llus
tam
bu
sisi
Pa
rasi
ro c
oif
fait
i
Fa
ng
en
sis
cave
rna
rum
Pu
rce
llia
illu
stra
ns
Ao
raki
tu
mid
ata
Pe
tta
lus
lam
pe
tid
es
Me
tag
ove
a p
hili
pi
Su
zuki
elu
s sa
ute
ri
Hu
ita
ca v
en
tra
lis
Ch
ileo
gov
ea
joca
sta
Ne
og
ove
a s
p
Ra
kaia
so
ren
sen
i d
igit
ata
Au
stro
pu
rce
llia
sco
pa
ria
Ka
rrip
urc
elli
a p
eck
oru
m
Pa
rap
urc
elli
a a
ma
tola
sp
nov
Pu
rce
llia
law
ren
cei s
p n
ov
Ch
ileo
go
vea
oe
dip
us
Sty
loce
llus
ram
bla
e
Ka
rrip
urc
elli
a s
ierw
ald
ae
Pu
rce
llia
tra
nsv
aa
lica
Pa
rap
urc
elli
a m
inu
ta s
p n
ov
Pa
rap
urc
elli
a s
tare
ga
i sp
nov
Pe
tta
lus
cim
icifo
rmis
Pa
rap
urc
elli
a f
issa
Fa
ng
en
sis
spe
lae
us
Tro
glo
siro
ae
llen
i
Pa
rap
urc
elli
a s
ilvic
ola
Sty
loce
llus
tam
bu
sisi
Pa
rap
urc
elli
a m
on
tico
la
Pa
rap
urc
elli
a n
ata
lia s
p n
ov
Ao
raki
cry
pta
Ra
kaia
ma
gn
a a
ust
ralis
Pu
rce
llia
gri
swo
ldi
sp
no
v
Sp
ele
osi
ro a
rga
sifo
rmis
Au
stro
pu
rce
llia
wo
od
wa
rdi
Tro
glo
siro
lo
ng
ifoss
a
Pa
rap
urc
elli
a c
onv
exa
sp
nov
Sir
o r
ub
en
s
Pa
rap
urc
elli
a p
ere
gri
na
tor
Cyp
ho
ph
tha
lmu
s d
uri
cori
us
Pa
rap
urc
elli
a r
um
pia
na
Ra
kaia
so
lita
ria
Ra
kaia
iso
lata
Ao
raki
lo
ng
ita
rsa
Ra
kaia
lin
dsa
yi
Ra
kaia
me
dia
Ao
raki
in
erm
a
Ao
raki
de
nti
cula
Sir
o a
caro
ide
s
Tro
glo
siro
nin
qu
a Ka
rrip
urc
elli
a h
arv
eyi
99 51 69
75 92 95 8581
80 69
94
6872
9998
74 83
86 86 5665
6379
8877 100
9381
81 87
68
97
Fig12
(A)Navajorugs
summarisingresults
forcladesof
interestanalysed
underparsim
onyusingcombineddiscreteandcontinuous
dataT
hesearch
conditionsvaried
acrossequalw
eights(EW)and
impliedweightswith
kvalues
rang
ingfrom
1to6andtherelativ
eweigh
tofthe
continuous
datasettothediscretedatasetranging
from
81
to18(B)S
trictconsensus
ofalltrees
foun
dinwhich
thegenus
Parap
urcelliaisfoun
dtobe
aderivedgrou
pwith
inthePettalid
ae(and
mon
ophyletic)Thiscorrespon
dstoalltreesun
derE
Wk=4k=5andk=6andk=3fordatapartition
weightin
gratio
sof8
11
11
2
14
and18(C)S
trictcon
sensus
ofalltreesinwhich
thegenusParapurcelliaisfoun
dtobe
basalw
ithinthePettalid
ae(and
paraphyletic)Thiscorrespo
ndstoalltreesun
derk
=1andk=2andk=3ford
ata
partition
weightin
gratio
sof41
and21N
umberson
thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom
analysesofthe11
weightedcombineddatasetunderEWk
=1and
k=6respectiv
ely
398 Invertebrate Systematics B L de Bivort and G Giribet
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)
The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies
We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae
Position of Parapurcellia peregrinator
Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses
Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal
P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae
Biogeography of the South African pettalids
In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons
One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)
Note added in proof
During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description
Systematic revision of South African pettalids Invertebrate Systematics 399
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Acknowledgements
We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871
References
Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x
Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x
Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x
Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x
Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046
Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044
Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x
de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029
de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x
Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391
Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119
Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211
Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354
Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76
Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2
Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014
Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830
Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16
Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2
Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)
Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18
Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x
Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x
Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x
Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x
Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087
Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073
Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312
Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)
Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)
Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)
Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280
400 Invertebrate Systematics B L de Bivort and G Giribet
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208
Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400
Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92
Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871
Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231
Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140
Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)
Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2
Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)
Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508
Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241
Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243
Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304
Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121
Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252
Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x
Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390
Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13
Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)
Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105
Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151
Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023
Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60
Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331
Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242
Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6
Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78
Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90
Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188
Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186
StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56
Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508
Manuscript received 23 May 2010 accepted 23 September 2010
Systematic revision of South African pettalids Invertebrate Systematics 401
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Appendix 1 List of material examined by light microscopy to generate the continuous data set
Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand
402 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)
1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its
surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation
5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and
Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia
12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)
13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or
unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca
ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit
19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor
bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral
process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented
centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally
26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp
article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article
28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles
29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this
dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference
31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 403
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Appendix 2 (continued )
32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus
ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)
36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip
(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort
of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and
anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully
visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings
45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis
(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface
(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal
plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)
56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins
appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families
58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua
59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata
60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail
61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)
404 Invertebrate Systematics B L de Bivort and G Giribet
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de
Bivort et al (2010)
1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal
opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the
maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width
21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the
length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent
35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities
of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum
(continued next page )
Systematic revision of South African pettalids Invertebrate Systematics 405
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis
Appendix 3 (continued )
52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the
length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum
54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point
(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV
divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the
scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and
the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal
thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody
74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer
75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the
chelicer is unornamented
406 Invertebrate Systematics B L de Bivort and G Giribet
httpwwwpublishcsiroaujournalsis