Undergraduate thesis in biology

Opisthobranchia: A taxonomic and biological review with emphasis on the families Chromodorididae and

Phyllidiidae together with field notes from South East Sulawesi, Indonesia

Lars Karlsson

LiU-IFM-Biol-Ex-892

Avdelning, Institution Division, Department

Avdelningen för biologi Institutionen för Fysik och mätteknik

Datum Date 010330

Språk Language

Rapporttyp Report category

ISBN LiU-IFM-Biol-Ex-892

Engelska/English x Examensarbete x D-uppsats

ISRN LiU-Biol-Ex-350

Handledare: Per Milberg Institution: Biology Department, IFM, University of Linköping

URL för elektronisk version

Titel Title: Opisthobranchia: A taxonomic and biological review with emphasis on the families Chromodorididae and Phyllidiidae together with field notes from South East Sulawesi, Indonesia Författare Author: Lars Karlsson

Sammanfattning Abstract This study gives an overview regarding Opisthobranchia (Mollusca, Gastropoda) and its taxonomy and some important biological facts. It also contains two parts of information regarding fieldwork conducted in Indonesia (Sulawesi) in cooperation with “Operation Wallacea” (a joint venture between a British company and an Indonesian research institute). The first of the two parts regarding fieldwork, gives a checklist of “Opisthobranchs of the Wakatobi Marine National Park” containing a total of 297 identified or suspected (provisionally identified) species (1997-1999). In the survey season 1999: 3523 opisthobranch observations were recorded, belonging to 235 species (identified or suspected) and 73 identified genera. During the whole survey 60 % of the opisthobranchs were identified to species level. The largest order was the Doridina, both in species number and number of genera. Fourteen percent of the opisthobranch species found in 1999 made up seventy-five percent of the total number of observations recorded. The second of the two parts regarding fieldwork is a description and analysis of spatial distribution of nudibranchs according to benthic structure (especially the families: Chromodorididae and Phyllidiidae since they are the most abundant). This survey was conducted at two sites, at two depths (around 3 and 12 m) and at two times a day (“am/pm”). During this survey 252 nudibranchs belonging to at least 19 species were recorded. According to a Detrended Correpondance Analysis, species composition seems to vary with benthic forms and depth.

Nyckelord Keywords: Opisthobranchia, Nudibranchia, Chromodorididae, Phyllidiidae, Operation Wallacea, Sulawesi, opisthobranch surveys, spatial distribution

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Contents Contents 1 Abstract 2 1 Review of opisthobranch taxonomy & biology 3 1.1 Introduction 3 1.2 Taxonomy of Gastropoda/Opisthobranchia 3 1.3 Morphology, ecology and physiology in Opisthobranchia 6 1.4 Nudibranchia 11 1.5 Chromodorididae 15 1.6 Phyllidiidae 18 1.7 References 18 2 Checklist of opisthobranchs of the Wakatobi National Park, SE Sulawesi, Indonesia 21 2.1 Introduction 21 2.2 Methods 21 2.3 Results 23 2.4 Discussion 26 2.5 References 28 3 Spatial distribution of nudibranchs according to benthic structure 29 3.1 Introduction 29 3.2 Material and methods 29 3.3 Results 31 3.4 Discussion 35 3.5 References 36 4 Conclusions and recommendations 37 5 Acknowledgements 38 Plates A B Appendices I II

2

Opisthobranchia: A taxonomic and biological review with emphasis on the families Chromodorididae and Phyllidiidae together with field

notes from South East Sulawesi, Indonesia

Lars Karlsson, Biology Department, IFM, University of Linköping, 2001.

Abstract

This study gives an overview regarding Opisthobranchia (Mollusca, Gastropoda) and its taxonomy and some important biological facts. It also contains two parts of information regarding fieldwork conducted in Indonesia (Sulawesi) in cooperation with “Operation Wallacea” (a joint venture between a British company and an Indonesian research institute). The first of the two parts regarding fieldwork, gives a checklist of “Opisthobranchs of the Wakatobi Marine National Park” containing a total of 297 identified or suspected (provisionally identified) species (1997-1999). In the survey season 1999: 3523 opisthobranch observations were recorded, belonging to 235 species (identified or suspected) and 73 identified genera. During the whole survey 60 % of the opisthobranchs were identified to species level. The largest order was the Doridina, both in species number and number of genera. Fourteen percent of the opisthobranch species found in 1999 made up seventy-five percent of the total number of observations recorded. The second of the two parts regarding fieldwork is a description and analysis of spatial distribution of nudibranchs according to benthic structure (especially the families: Chromodorididae and Phyllidiidae since they are the most abundant). This survey was conducted at two sites, at two depths (around 3 and 12 m) and at two times a day (“am/pm”). During this survey 252 nudibranchs belonging to at least 19 species were recorded. According to a Detrended Correpondance Analysis, species composition seems to vary with benthic forms and depth.

3

1 Review of opisthobranch taxonomy & biology 1.1 Introduction Opisthobranchs or “sea slugs” are a large, exclusively (except for the freshwater Ancylodoris baicalensis) marine group within the Gastropoda (Wägele & Willan, 2000). They are often colourful and display several forms of crypsis, special resemblance, aposematic coloration and mimicry (Gosliner & Behrens, 1990; Tullrot, 1998). Most opisthobranchs lack the protection of a shell as adults (some of them do have a shell but it is not functioning as a protective device since it is reduced and sometimes also internal). Instead these organisms rely on other defences. Some opisthobranchs use spicules as an alternative to shell, others are cryptic, some utilize the nematocysts from their prey and a few use autotomy. One common defensive systems, though, is chemical defence (Faulkner & Ireland, 1977; Todd, 1981; Faulkner & Ghiselin, 1983; Gunthorpe & Cameron, 1987; Avila, 1995; Cimino et al., 1999). This is the case for both Chromodorididae and Phyllidiidae (Nudibranchia). Both groups prey on sponges, which often contain bad tasting or even toxic compounds. The opisthobranchs then accumulate (or convert) some of these compounds and thereby become less interesting for potential predators. This habit has made them interesting for pharmaceutical screening as many of the compounds they use are suspected to have pharmaceutical value (Bohlin, 1989). Another interesting feature is that the chemical toxicity of these species often is coupled with strong examples of aposematism. Often, the same type of strong warning coloration is used by many different opisthobranchs. If the opisthobranchs with the similar coloration are all unpalatable, then it is an example of Müllerian mimicry. On the other hand, if there are palatable species among the ones with similar coloration, then it is an example of Batesian mimicry (Ruppert & Barnes, 1994). One interesting discovery is that there is also a flatworm species, which seems to mimic a phyllidiid nudibranch in order to avoid predation (Newman et al., 1994). 1.2 Taxonomy of Gastropoda/Opisthobranchia There are many different and confusing views on Gastropod systematics. In most publications, the class Gastropoda consists of three subclasses: Prosobranchia, Pulmonata and Opisthobranchia (Ruppert & Barnes, 1994; Hayward & Ryland, 1995; Thompson, 1976). This class is not considered by some authors to reflect the phylogeny of gastropods and a newer (cladistic) classification divides the Gastropoda into two clades: Eogastropoda and Orthogastropoda. In the latter group is a clade called Apogastropoda, which contain Heterobranchia and Caenogastropoda. The former group, in turn, consists of three clades: Heterostropha, Opisthobranchia and Pulmonata (Ponder & Lindberg, 1997). Most authors agree that within the Opisthobranchia there are at least eight groups, although they have been given different taxonomic ranks. These subgroups within the Opisthobranchia are (Marshall & Willan 1999): Cephalaspidea (Bubble shells), Anaspidea (Sea hares), Notaspidea (Side-gilled slugs), Acochlidiacea (Caddis-slugs), Sacoglossa (Sap-sucking sea slugs), Thecosomata (Sea butterflies), Gymnosomata (Naked sea butterflies) and Nudibranchia (Nudibranchs). Sometimes other groups are included in the Opisthobranchia, for example Pyramidellacea (Ruppert & Barnes, 1994) Pyramidellidae (Hayward & Ryland, 1995), Entomotaeniata (Willan & Coleman, 1984), Rhodopida (Hansson, 1998) and Rhodopemorpha (Beesley et al., 1998). Many of these groups have previously been assigned different names and some of these synonyms are shown in Table 1 together with examples of different views regarding levels of taxa.

4

Table 1. Systematic review of Opisthobranchia according to different authors. Marshall & Willan,

1999 Rudman & Willan, 1998

Hansson, 1998 Hayward & Ryland, 1995

Ruppert & Barnes,

(In Beesley et al., 1998)

1994 *

Regnum Animalia Animalia Phylum Mollusca Mollusca Mollusca Subphylum Conchifera Superclassis Classis Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Subclassis Opisthobranchia Opisthobranchia Heterobranchia Opisthobranchia Opisthobranchia Infraclassis Euthyneura Superordo Opisthobranchia Ordo Cephalaspidea Cephalaspidea = Cephalaspid(e)a = Cephalaspidea Cephalaspidea Bullomorpha Bullomorpha = Tectibranchida Ordo Anaspidea Anaspidea = Anaspid(e)a = Anaspidea Anaspidea = Aplysiomorpha Aplysiomorphi(id)a Aplysiacea Ordo Notaspidea Notaspidea = Notaspidea Notaspidea Pleurobranchomorpha Ordo Acochlidia Acochlidea = Acochlidiomorph(id)a = Acochlidiacea Acochlidioidea Acochlidia Acochlidioid(e)a Ordo Sacoglossa Sacoglossa = Sacogloss(id)a = Sacoglossa Sacoglossa Ascoglossa = Ascogloss(id)a Monostichoglossa = Pellibranchiata Ordo Thecosomata Thecosomata Thecosomat(id)a Thecosomata Thecosomata Ordo Gymnosomata Gymnosomata Gymnosomat(id)a Gymnosomata Gymnosomata Ordo Rhodopemorpha Rhodopida Pyramidellidae Pyramidellacea Ordo Nudibranchia Nudibranchia = Nudibranchi(d)a Nudibranchia Nudibranchia Behrens, 1991 and Thompson, 1988 ** Willan & Coleman, Thompson, 1976 Gosliner, 1987 1984 Regnum Phylum Mollusca Subphylum Superclassis Classis Gastropoda Gastropoda Gastropoda Subclassis Opisthobranchia Opisthobranchia Opisthobranchia Opisthobranchia Infraclassis Superordo Ordo Cephalaspidea = Bullomorpha Cephalaspidea Bullomorpha Bullomorpha Ordo Anaspidea = Aplysiomorpha Anaspidea Aplysiomorpha Aplysiomorpha Ordo Notaspidea = Pleurobranchomorpha Notaspidea Pleurobranchomorpha Pleurobranchomorpha Ordo Acochlidiacea Acochlidiacea Acochlidiacea Ordo Sacoglossa = Sacoglosssa Sacoglossa Sacoglosssa Ascoglossa Ordo Thecosomata Thecosomata Ordo Gymnosomata Gymnosomata Ordo Entomotaeniata Ordo Nudibranchia Nudibranchia Nudibranchia Nudibranchia * Note: Thecosomata and Gymnosomata are sometimes called: "pteropods". ** Note: Excluding the pteropods and the pyramidellids. Going further down in the classification, a summary of opisthobranch families is presented in Table 2. (This classification is also used by a very valuable database on the Internet, “The Sea Slug Forum” (Rudman, 2000)).

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Table 2. Summary of ophistobranch families (after Beesley et al, 1998). OPISTHOBRANCHIA Order CEPHALASPIDEA Superfamily ACTEONOIDEA Family Acteonidae Family BuIlinidae Family Hydatinidae Superfamily RINGICULOIDEA Family Ringiculidae Superfamily CYLINDROBULLOIDEA Family Cylindrobullidae Superfamily DIAPHANOIDEA Family Diaphanidae Superfamily PHILINOIDEA Family Cylichnidae Family Retusidae Family Philinidae Family Philinoglossidae Family Aglajidae Family Gastropteridae Superfamily HAMINOEOIDEA Family Haminoeidae Family Bullactidae Family Smaragdinellidae Family Incertae sedis Superfamily BULLOIDEA Family Bullidae Superfamily RUNCINOIDEA Family Runcinidae Family Ilbiidae Superfamily INCERTAE SEDlS Family Notodiaphanidae Order ACOCHLIDEA Superfamily ACOCHLIDIOIDEA Family Acochlidiidae Family Hedylopsidae Superfamily MICROHEDYLOIDEA Family Asperspinidae Family Microhedylidae Family Ganatidae Order RHODOPEMORPHA Family Rhodopidae

Order SACOGLOSSA Superfamily OXYNOOIDEA Family Volvatellidae Family Oxynoidae Family Juliidae Superfamily ELYSIOIDEA Family Plakobranchidae Family Elysiidae Family Boselliidae Family GascoigneIlidae Family Platyhedylidae Superfamily LIMAPONTIOIDEA Family Caliphyllidae Family Costasiellidae Family Hermaeidae Family Limapontiidae Order ANASPIDEA Superfamily AKEROIDEA Family Akeridae Superfamily APLYSIOIDEA Family Aplysiidae Order NOTASPIDEA Superfamily TYLODINOIDEA Family Tylodinidae Family Umbraculidae Superfamily PLEUROBRANCHOIDEA Family Pleurobranchidae Order THECOSOMATA Family Limacinidae Family Cavoliniidae Family Peraclididae Family Cymbuliidae Family Desmopteridae Order GYMNOSOMATA Suborder GYMNOSOMATA Family Pneumodermatidae Family Notobranchaeidae Family Cliopsidae Family Clionidae Suborder GYMNOPTERA Family Hydromylidae

Order NUDIBRANCHA Suborder DORIDINA Superfamily ANADORIDOIDEA Family Corambidae Family Goniodorididae Family Onchidorididae Family Polyceridae Family Gymnodorididae Family Aegiretidae Family Vayssiereidae Superfamily EUDORIDOIDEA Family Hexabranchidae Family Dorididae Family Chromodorididae Family Dendrodorididae Family Phyllidiidae Suborder DENDRONOTINA Family Tritoniidae Family Bornellidae Family Marianinidae Family Hancockiidae Family Dotidae Family Scyllaeidae Family Tethydidae Family Phylliroidae Family Lomanotidae Suborder ARMININA Family Arminidae Family Doridomorphidae Family Charcotiidae Family MadreIlidae Family Zephyrinidae Family Pinufiidae Suborder AEOLIDINA Family Flabellinidae Family Eubranchidae Family Aeolidiidae Family Glaucidae Family Embletoniidae Family Tergipedidae Family Fionidae

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1.3 Morphology, ecology and physiology in Opisthobranchia Figure 1 shows a broad characterisation of grades of organisation in the opisthobranchs. The circle “1” is considered to represent the oldest and most “primitive” level.

Figure 1. Grades (time-development level) of organisation in the opisthobranchs (Beesley et al, 1998) (Reproduced by special permission from Richard Willan and Australian Biological Resources Study, Environment Australia) As can be seen, within each order there are usually several grades of organisation. In spite of this, it is also possible to see that the different groups of nudibranchs are considered to have reached the “highest” forms of organisation (Beesley et al., 1998). There is a strong coupling between organisation grades and morphological appearances, as for instance, when the shell is reduced or lacking completely. The first way of identifying opisthobranch orders is usually by external examination as the morphology often gives clues to which order a specimen can be placed. Figure 2-9 shows some typical members of eight opisthobranch groups.

a b Figure 2. (a) Cephalaspidea, Bulla ampulla. (b) Cephalaspidea, Chelidonura sandrana (Thompson, 1976). (Thompson, 1976).

7

Figure 3. Anaspidea, Aplysia depilans Figure 4. Notaspidea, Pleurobranchus (Thompson, 1976). membranaceus (Thompson, 1976).

a b Figure 5. (a) Acochlidiacea, Hedylopsis loricata (b) Acochlidiacea, Microhedyle sp (Thompson, 1976). (Thompson, 1976).

a b

Figure 6. (a) Sacoglossa, Stiliger fuscatus (b) Sacoglossa, Elysia marginata (Thompson, 1976). (Thompson, 1976).

8

a b Figure 7. (a) Thecosomata, Spirellatidae (b) Thecosomata, Peraclidae. (Thompson, 1976). (Thompson, 1976).

a b Figure 8. (a) Gymnosomata, Crucibranchaea (b) Gymnosomata, Clione limacina. macrochira (Thompson, 1976). (Thompson, 1976).

a b Figure 9. (a) Nudibranchia, Polycera capensis (b) Nudibranchia, Tritonia festiva (Thompson, 1976). (Thompson, 1976).

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In most books and surveys only five orders are treated. Usually Acochlidiacea is not treated since they are minute and difficult to collect (requires sieving through sand). Thecosomata and Gymnosomata are also difficult to collect since they are pelagic. Accordingly, most estimates of opisthobranch species diversity are based on just five groups. One exception to this is Thompson (1988) who states that there is “about 3000 species” world-wide. (For more information regarding opisthobranch diversity, see Table 3.) Table 3. Review of opisthobranch diversity in chosen areas. Opisthobranchia (A = Estimations, B = Found species)

A Area Source Remarks: 3000 Worldwide Thompson, 1976 700 Tropical Western

Pacific Marshall & Willan, 1999 Not including: Acochlidea, Thecosomata, Gymnosomata

600 Pacific "Equatorial Belt"

Marshall & Willan, 1999 Not including: Acochlidea, Thecosomata, Gymnosomata

550 Great Barrier Reef

Marshall & Willan, 1999 Not including: Acochlidea, Thecosomata, Gymnosomata

350 S. Barrier Reef Marshall & Willan, 1999 Not including: Acochlidea, Thecosomata, Gymnosomata 120 Sweden Hansson, 1998 Eight orders B Area Source Remarks:

561 Madang (PNG) Gosliner (in M & W, 1999) Not including: Acochlidea, Thecosomata, Gymnosomata 462 Great Barrier

Reef Marshall & Willan, 1999 Checklist of Great Barrier Reef Opisthobranchia (eight

orders) 439 Guam Carlson & Hoff (in M & W,

1999) Not including: Acochlidea, Thecosomata, Gymnosomata

390 South Africa Gosliner, 1987 Not including Acochlidea 389 Marshall Islands Johnson & Boucher (in M

& W, 1999) Not including: Acochlidea, Thecosomata, Gymnosomata

240 Suva (Fiji) Brodie & Brodie ( in M & W, 1999)

Not including: Acochlidea, Thecosomata, Gymnosomata

262 Heron Island Marshall & Willan, 1999 Not including: Acochlidea, Thecosomata, Gymnosomata 178 Norway Moen & Svensen, 1999 156 British Isles Thompson, 1988 Not including: Thecosomata, Gymnosomata 105 New Caledonia Risbec, 1928 (in M & W,

1999) Not including: Acochlidea, Thecosomata, Gymnosomata

All opisthobranchs are hermaphrodite and copulation is normally reciprocal (Thompson, 1988; Hayward & Ryland, 1995). The reproductive organs are present in all adults on the right-hand side of the body (Hayward & Ryland, 1995). Most species spawn eggs, which hatch as free swimming, shelled, veliger larvae (Thompson, 1988; Hayward & Ryland, 1995). In Figure 10 there are some examples of typical spawn masses and in Figure 11 there are examples of veliger larvae.

10

Figure 10. Examples of opisthobranch spawn masses (Behrens, 1991).

11

Figure 11. Examples of opisthobranch veliger larva (Thompson, 1988). The shelled veliger larva is normally planktotrophic while those species that hatch as miniature adults are normally lecitotrophic. There is a greater proportion of these species in polar seas (Willan & Coleman, 1984) Internally, the opisthobranchs display detorsion and many are secondarily bilaterally symmetrical. They have one auricle and one nephridium (Ruppert & Barnes, 1994). Most opisthobranchs are carnivorous, although sacoglossans and aplysiids are herbivorous (Hayward & Ryland, 1995). 1.4 Nudibranchia The largest group within the Opisthobranchia is the “Nudibranchia”. Currently, there are conflicting views whether Nudibranchia is monophyletic (Wägele & Willan, 2000) or polyphyletic (Tholleson, 1998). Wägele & Willan (2000) goes even further and propose a new taxon “Nudipleura” as a monophyletic group, which contains Pleurobranchoidea and Nudibranchia. As with the opisthobranchs, there are different views on nudibranch systematic and a short review is presented in Table 4. (A historical summary of the nudibranch divisions is shown in Table 5.)

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Tab

le 4

. Sys

tem

atic

revi

ew o

f Nud

ibra

nchi

a ac

cord

ing

to v

ario

us a

utho

rs.

Wäg

ele

& W

illan

, 20

00 C

lade

s!

M

arsh

all &

Will

an,

1999

R

udm

an &

Will

an,

1998

(In

Bee

sley

et

al.,

1998

)

Han

sson

, 199

8 D

ebel

ius,

199

6 Th

omps

on, 1

988

Wel

ls &

Bry

ce,

1993

B

ehre

ns, 1

991

Gos

liner

, 198

7 W

illan

& C

olem

an,

1984

Anth

obra

nchi

a Su

bord

o

Dor

idin

a An

thob

ranc

hi(n

)a =

D

orid

acea

D

orid

oide

a D

orid

acea

An

thob

ranc

hia

* C

lado

bran

chia

Dor

id(a

c)in

a *

Su

bord

o

Den

dron

otin

a C

lado

bran

chin

a #

Den

dron

otac

ea

Den

dron

otoi

dea

Den

dron

otac

ea

Cla

dobr

anch

ia #

Su

bord

o

Arm

inin

a N

otas

pide

(in)a

=

Arm

inac

ea

Arm

inoi

dea

Arm

inac

ea

Pl

euro

bran

chom

orph

(in)a

Subo

rdo

Ae

olid

ina

Ae

olid

acea

Ae

olid

oide

a

Aeol

idac

ea

Infr

aord

o

Supe

rfam

ilia

Dor

idoi

dea

* E

udor

idoi

dea

* D

orid

acea

D

endr

onot

oide

a

* Pol

ycer

oide

a

# D

endr

onot

acea

Ar

min

oide

a

* Ana

dorid

oide

a

# Ar

min

acea

Ae

olid

oide

a

* Nam

e?

#

Aeol

idac

ea

* Nam

e?

* Nam

e?

# D

endr

onot

oide

a

#

Arm

inoi

dea

# Ae

olid

oide

a

Fam

ilia

Subf

amili

a Tr

ibus

G

enus

Su

bgen

us

Spec

ies

Guv

erne

d by

ICZN

G

uver

ned

by IC

ZN

Guv

erne

d by

ICZN

G

uver

ned

by IC

ZN

Guv

erne

d by

ICZN

G

uver

ned

by IC

ZN

Guv

erne

d by

ICZN

Not

e: N

ew ta

xon

prop

osed

"N

udip

leur

a" =

N

udib

ranc

hia

+ Pl

euro

bran

choi

dea

Not

e: N

udib

ranc

hia

= An

thob

ranc

hia

(=

Euct

enid

ieac

ea =

C

teni

diac

ea) +

C

lado

bran

chia

(=

Acte

nidi

acea

)

Not

e: In

Dor

idoi

dea

all s

peci

es a

re

holo

hepa

tic.

Not

e: D

orid

acea

is

divi

ded

in th

ree

grou

ps:

cryp

tobr

anch

s,

poro

stom

es a

nd

phan

erob

ranc

hs

Not

e: S

omet

imes

th

e An

thob

ranc

hia

is d

ivid

ed in

to

cryp

tobr

anch

s an

d ph

aner

obra

nchs

Foot

note

; IC

ZN =

Inte

rnat

iona

l Cod

e of

Zoo

logi

cal N

omen

clat

ure

(Fou

rth E

ditio

n 20

00)

13

Table 5. Historical summary of the nudibranch divisions (Wägele & Willan, 2000).

The most difficult nudibranch group to characterise is the Arminina/Arminacea/Arminoidea group (Figure 12).

a b Figure 12. (a) Arminina, Armina loveni (b) Arminina, Dirona albolineata (Todd, 1981). (Thompson, 1976). The other nudibranch groups are somewhat easier to describe as seen in Figures 13-15.

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a b Figure 13. (a) Doridina, Ceratosoma cornigerum (b) Doridina, Plocamopherus ceylonicus (Thompson, 1976). (Thompson, 1976).

a b Figure 14. (a) Dendronotina, Doto yongei (b) Dendronotina, Hancockia burni (Thompson, 1976). (Thompson, 1976).

a b Figure 15. (a) Aeolidina, Pseudovermis mortoni (b) Aeolidina, Facelina annulicornis (Thompson, 1976). (Thompson, 1976).

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As with the opisthobranchs, the number of nudibranch species is largest in the tropics (Willan & Coleman, 1984). As most recent publications treat Nudibranchia as an order or a suborder, the corresponding lower levels are either suborders or (super) families. Regardless, Nudibranchia is usually considered to contain four subgroups: Doridina/Doridacea/Dorodoidea, Aeolidina/Aeolidacea/Aeolidoidea, Arminina/Arminacea/Arminoidea and Dendronotina/Dendronotacea/Dendronotoidea. The group with the largest number of species is the Doridina/Doridacea/Dorodoidea. In “Checklist of Great Barrier Reef Opisthobranchia” (Marshall & Willan, 1999) 281 nudibranch species are recorded totally and out of them, 204 species belong to the Doridina/Doridacea/Dorodoidea group. According to Wägele & Willan (2000) Nudibranchia consists of about 190 genera and 60 families. In Table 6, examples of various nudibranch estimations or records are shown. Table 6. Review of nudibranch diversity in chosen areas.

Nudibranchia (A = estimations, B = found species) A Area Source Remarks

3000 Worldwide Willan & Coleman 1984 800 Tropical Indo Pacific Willan & Coleman 1984 B Area Source Remarks

281 Great Barrier Reef Marshall & Willan, 1999 Checklist of Great Barrier Reef Opisthobranchia

269 Australia Willan & Coleman 1984 Checklist of Australian Nudibranchia 250 Mediterreanean Cattaneo-Vietti et al, 1990 Checklist of the Mediterranean

Nudibranchs 241 South Africa Gosliner, 1987 Southern African Opisthobranch

Species 108 British isles Picton & Morrow, 1994 British Check List 108 British Isles Thompson, 1988 British Check List 80 Sweden Hansson, 1998 Included 2 species of Notaspidea!

The life span of different nudibranch species varies between a few weeks and two years. Usually three ecological groupings are used: “Subannual“, “Annual”, and “Biennial” species. The subannual species are normally feeding on seasonally variable resources while the biennial species are feeding on stable diets. All nudibranchs appear to be semelparous (Todd, 1981). The majority of anthobranchs eat sponges or bryozoans and most cladobranchs eat coelenterates. Other nudibranch diets include ascidians, barnacles, crustaceans, and even the spawn of other opisthobranchs (Willan & Coleman, 1984). The most complete review regarding nudibranch food has been provided by McDonald & Nybakken (1997). Two families of the Doridina/Doridacea/Dorodoidea group are very common both in species number and abundance in the Indo-Pacific: Chromodorididae and Phyllidiidae. 1.5 Chromodorididae This family has a lot of species (more than 300 according to Wells & Bryce, 1993) and many are difficult to distinguish. For instance, in order to separate the genus Chromodoris and the genus Hypselodoris, the researcher usually has to dissect, prepare and compare the radula (Figure 16), which is present in all chromodorids. This takes a lot of skill and access to a good microscope or even a scanning electron microscope (SEM).

16

Figure 16. Examples of opistobranch radular teeth (Photos taken by Terry Gosliner, in Behrens 1991). Luckily, after this has been done it is often possible to use minor differences in external form and colour to identify many species. One key characteristic is the shape and form of the rhinophores (Figure 17).

17

Figure 17. Various shapes and forms of ophistobranch rhinophores (Behrens, 1991). Especially within the Chromodorididae there are many “colour groups” which are very similar in appearances (Rudman, 1973, 1977, 1982, 1984, 1991; Gosliner & Johnson 1999). Often their appearance is quite colourful and it is thought that this is an example of aposematism. As several species then share the same general colour markings it is suspected that this is also an example of Müllerian mimicry and that the species take advantage of similar coloration in order to avoid predation. The deterrent, which makes this work, is normally a substance acquired from sponges, which are the normal food for chromodorids. In some cases it is just a question of accumulating useful compounds from the sponges but there are also some chromodorids which can convert ingested substances to a more “toxic” state (Karuso, 1987).

18

1.6 Phyllidiidae Another interesting family is the Phyllidiidae, it is also a member of the Doridina/Doridacea/Dorodoidea group and according to Wells & Bryce (1993) the family consists of about 70 species. The phyllidiids lack a radula (Brunckhorst, 1993; Yonow, 1996). Instead they “suck” in pieces of sponges which they utilize as food. Together with the dendrodorids (which also lacks radula) the phyllidiids constitutes a group which is called “porostomata”. The phyllidiids can secrete a “milky” fluid when handled and this fluid can work as a poison. This was first noted when a phyllidiid accidentally killed all the lobsters in an aquarium where it was put for storage. After this discovery a lot of research have been conducted on chemical substances in different species of phyllidiids (Karuso, 1987). (Some chemical structures of these compounds are shown in Figure 18-19).

Figure 18. Toxin from Phyllidia varicosa, Figure 19. Major metabolite from 9-isocyanopupukeananae P. pulitzeri, axisonitrile-1 (Karuso, 1987). and its 2-isomer (Karuso, 1987). The phyllidiids are mainly restricted to tropical areas and there are just a few species of this family in more temperate water (Gosliner & Behrens, 1988). 1.7 References Avila, C. 1995. Natural products of opisthobranch molluscs: a biological review.

Oceanography and Marine Biology: an Annual Review 33: 487-559. Beesley, P. L., Ross, G. J. B. & Wells, A. (eds). 1998. Mollusca: The Southern Synthesis.

Fauna of Australia. Vol. 5. CSIRO Publishing. Melbourne. Behrens, D. W. 1991. Pacific Coast nudibranchs: a guide to the opisthobranchs, Alaska to

Baja California. Seachallenger. Monterey. Bohlin, L. 1989. Pharmacologically active compounds from marine organisms. Acta

Pharmacetica Nordica 1: 175-182. Brunckhorst, D. J. 1993. The systematics and phylogeny of phyllidiid nudibranchs

(Doridoidea). Australian Museum. Sydney. Cattaneo-Vietti, R., Chemello, R. and Gianuzzi-Savelli, R. 1990. Atlas of Mediterranean

nudibranchs. La Conchiglia. Roma. Cimino, G., Fontana, A. and Gavagnin, M. 1999. Marine Opisthobranch Molluscs:

Chemistry and Ecology in Sacoglossans and Dorids. Current Organic Chemistry 3: 327-372. Debelius, H. 1996. Nudibranchs and sea snails Indo Pacific field guide. IKAN –

Unterwasserarchiv. Frankfurt. Faulkner, D. J. and Ireland, C. 1977. The chemistry of some opisthobranch molluscs. Pp 23-

34 in D. J. Faulkner and W. H. Fenical (eds). Marine natural products chemistry. Plenum Press. New York.

19

Faulkner, D. J. and Ghiselin, M. T. 1983. Chemical defence and evolutionary ecology of dorid nudibranchs and some other opisthobranch gastropods. Marine Ecology – Progress Series 13: 295-301.

Gosliner, T. 1987. Nudibranchs of Southern Africa. Seachallenger. Monterey. Gosliner, T. M. and Behrens, D. W. 1988. A Review of the Generic Divisions Within the

Phyllidiidae with the Description of a New Species of Phyllidiopsis (Nudibranchia: Phyllidiidae) from the Pacific Coast of North America. The Veliger 30: 305-314.

Gosliner, T. M. and Behrens, D. W. 1990. Special Resemblance, Aposematic Coloration and Mimicry in Opisthobranch Gastropods. Pp 127-138 in M. Wicksten (ed). Symposium on the Adaptive Significance of Colour in Invertebrates. University Press Texas A & M.

Gosliner, T. M. and Johnson, R. F. 1999. Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with a review of the monophyletic clade of Indo Pacific species, including descriptions of twelve new species. Zoological Journal of the Linnean Society 125: 1-114.

Gunthorpe, L. and Cameron, A. M. 1987. Bioactive properties of extracts from Australian dorid nudibranchs. Marine Biology 94: 39-43.

Hansson, H. G. 1998. Sydskandinaviska marina flercelliga evertebrater, utgåva 2. Länsstyrelsen Västra Götaland. Göteborg.

Hayward, P. J. & Ryland, J. S. (eds). 1995. Handbook of the Marine Fauna of North – West Europe. Oxford University Press. Oxford, New York, Tokyo.

ICZN, Fourth Edition. 1999. International Code of Zoological Nomenclature. The International Trust for Zoological Nomenclature. London.

Karuso, P. 1987. Chemical Ecology of the Nudibranchs. Pp 32-60 in Scheuer, P. J. (ed). Bioorganic Marine Chemistry, Volume 1. Springer- Verlag. Berlin, Heidelberg.

McDonald, G. & Nybakken, J. 1997. A Worldwide Review of the food of Nudibranch Mollusks. The Veliger 40: 157-159.

Marshall, J. G. and Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef. Backhuys Publishers. Leiden.

Moen, F. E. & Svensen, E. 1999. Dyreliv i havet. KOM forlag, Kristiansund. Newman, L. J., Cannon, L. R. G. and Brunckhorst, D. J. 1994. A new flatworm

(Platyhelminthes: Polycladida) which mimics a Phyllidiid Nudibranch (Mollusca, Nudibranchia). Zoological Journal of the Linnean Society 110: 19-25.

Picton, B. E. & Morrow, C. C. 1994. A Field Guide to the Nudibranchs of the British Isles. Immel Publishing. London.

Ponder, W. F. and Lindberg, D. R. 1997. Towards a phylogeny of gastropod molluscs: an analysis using morphological characters. Zoological Journal of the Linnean Society 119: 83-265.

Rudman, W. B. 1973. Chromodorid opisthobranch Mollusca from the Indo-West Pacific. Zoological Journal of the Linnean Society 52: 175-199.

Rudman, W. B. 1977. Chromodorid opisthobranch Mollusca from East Africa and the tropical West Pacific. Zoological Journal of the Linnean Society 61: 351-397.

Rudman, W. B. 1982. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris quadricolour, C. lineolata and Hypselodoris nigrolineolata colour groups. Zoological Journal of the Linnean Society 76: 183-241.

Rudman, W. B. 1984. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo –West Pacific: a review of the genera. Zoological Journal of the Linnean Society 81: 115-273.

Rudman, W. B. 1991. Purpose in pattern: the evolution of colour in chromodorid nudibranchs. Journal of Molluscan Studies 57: 5-21.

Rudman, W. B. 2000. The Sea Slug Forum, www.austmus.gov.au/science/invert/mal/forum Ruppert, E. E. & Barnes, R. D. 1994. Invertebrate Zoology. Saunders College Publishing.

Fort Worth, Philadephia, San Diego.

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Thollesson, M. 1998. Nudibranch systematics and molecular data. PhD Thesis, Göteborgs Universitet.

Thompson, T. E. 1976. Biology of opisthobranch molluscs. Volume I. The Ray Society. London.

Thompson, T. E. 1988. Molluscs: Benthic Opisthobranchs. E. J. Brill / Dr W. Backhuys. Leiden, New York, København, Köln.

Todd, C. D. 1981. The ecology of nudibranch molluscs. Oceanography and Marine Biology: an Annual Review 19: 141-234.

Tullrot, A. 1998. Evolution of warning coloration in the nudibranch Polycera quadrilineata. PhD Thesis, Göteborgs Universitet.

Wells, F. E. & Bryce, C. W. 1993. Sea slugs of Western Australia. Western Australian Museum. Perth.

Willan, R. C. & Coleman, N. 1984. Nudibranchs of Australasia. Australasian Marine Photographic Index. Sydney.

Wägele, H. & Willan, R. C. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the Linnean Society 130: 83-181.

Yonow, N. 1996. Systematic revision of the family Phyllidiidae in the Indian Ocean province: Part 1 (Opisthobranchia: Nudibranchia: Dorodoidea). Journal of Conchology 35: 483-516.

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2 Checklist of opisthobranchs of the Wakatobi National Park, SE Sulawesi, Indonesia

2.1 Introduction Operation Wallacea (“Opwall”) is a joint venture between the Indonesian “Wallacea Development Institute” and a British company: Ecological Surveys Ltd. The co-operation started 1995 and the purpose was to promote conservation work in a remote Indonesian archipelago, Tukang Besi (Chart 1). The Operation Wallacea works with both land and marine projects, concentrating mainly on field surveys regarding diversity issues but it has also been involved in some projects promoting sustainable use of natural resources. This means, for instance, changing the fishing methods and promoting ecotourism in the area. A significant result of this work was achieved when “The Wakatobi Marine National Park” was created in July 1996. It is currently the second largest Indonesian “National Marine Park” (13000 km2) stretching between 05º15’S, 123º23’E and 06º08’S, 124º37’E (Stanzel & Newman, 1997). The name Wakatobi comes from the first letters of the four largest islands in the area (Wanci, Kaledupa, Tomea and Binongko). One aim of Operation Wallacea is to survey and publish checklists of the marine life in the national park. During 1997-99 a lot of work was geared towards a checklist of Opisthobranchia. This paper presents results from this work.

Chart 1. Tukang Besi archipelago, Indonesia (Expedia, 2001). 2.2 Methods The marine projects, run by Operation Wallacea, are concentrated around a small island, Hoga (Chart 2), situated near Kaledupa. During my visit 1999, two major projects were conducted, one regarding correlation between butterfly fish diversity and coral diversity and another one regarding opisthobranch diversity. Paying volunteers did nearly all work but there was also some scientific staff supervising the projects. The opisthobranch project started in 1997 (two

22

weeks) and continued for 10 months in 1998 and for 7 months in 1999. The main effort was done during 1999 when a lot of work was concentrated at opisthobranchs.

Chart 2. Hoga Island, Tukang Besi archipelago, Indonesia (Expedia, 2001). In the opisthobranch project, groups of diving volunteers concentrated on finding as many species of opisthobranchs as possible (down to a maximum depth of 30 m). For each recording they noted numbers of individuals, dates, depths and sizes. All species were photographed and unidentifiable species were also drawn by hand. The main identification books used at Hoga were: Allen & Steene, 1996; Behrens, 1991; Coleman, 1989; Debelius, 1996; Gosliner, 1987, Gosliner, et al, 1996; and Wells & Bryce, 1993. The collected data for 1999 (handwritten notes) were then taken to England and later also copied and sent to Sweden where the compilation (Appendix I) was made during the year of 2000. During this work it was possible to identify some more species with the help of the Internet, especially as the site “The Sea Slug Forum” proved to be very valuable (Rudman, 2000). At least one specimen of each species was conserved at Hoga for later shipment to the Bogor Zoological Museum. Also, from many species, a small tissue sample (large species) or a whole individual (small species) was taken and conserved. The aim with this material is to compare the DNA from the species and to do a cladistic analysis later on (Warren, 2000 and personal communication, October 1999*). Notes of the taxa found during 1997 or 1998 but not 1999 are added to “Appendix I” but these taxa are not included in calculations regarding “observations” or “occasions” due to lack of data. The reason for choosing the word “observations” instead of “individuals” is because each individual could have been recorded more than once. The term “occasions” is used since not all survey work was done diving. In a few cases surveys were made in the intertidal area during low tide. * Lindsay Warren, personal communication, October 1999 (Hoga Island).

Hoga Island

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2.3 Results A total of 60 different species were found during two weeks in September 1997. Between March and December 1998 another 115 opisthobranch species were found. The total species number for the whole period 1997-1999 is 297 species (Warren, 2000). During the season of 1999 (May-November), 3523 opisthobranchs were found during the survey. They belonged to a total of 235 identified or suspected (provisionally identified) species and to 73 identified genera (Appendix I). Table 1. Summary of “The 1999 Opisthobranch Survey – Operation Wallacea” (Indonesia) Anaspidea Cephalaspidea Notaspidea Nudibranchia Sacoglossa Species 9 36 5 161 23 Observations 116 316 42 2888 159

Two additional observations were of Colpodaspis thompsoni, which probably belong to Cephalaspidea but its current taxonomic status is uncertain (Marshall & Willan, 1999). The results regarding discovered species and genera for the periods 1997-1999, 1997-1998 and 1999 are shown in Figure 1 and 2.

Operation Wallacea

Anaspidea

Cephalaspidea

Notaspidea

Nudibranchia

Sacoglossa

0 50 100 150 200 250

Cat

egor

y

Number of species

1997-199919991997-1998

Figure 1. Summary of species, “Operation Wallacea - Opisthobranchia survey” 1997-1999 (Indonesia).

24

Operation Wallacea

Anaspidea

Cephalaspidea

Notaspidea

Nudibranchia

Sacoglossa

0 20 40 60

Cat

egor

y

Number of identified genera

1997-199919991997-1998

Figure 2. Summary of identified genera, “Operation Wallacea - Opisthobranchia survey” 1997-1999 (Indonesia). In Figure 3 there is a summary of the results of the identification effort.

Identification of Opisthobranchs 1997-99

Genus + speciesGenus Only higher taxa

Figure 3. Result of identification effort regarding 295 opisthobranch taxa, “Operation Wallacea - Opisthobranchia survey” 1997-1999 (Indonesia). The proportions (species and genera) between the nudibranch suborders for the period 1997-1999 are presented in Figure 4 and 5.

25

Nudibranchia 1997-99

AeolidinaArmininaDendronotinaDoridina

Figure 4. Proportion of species within nudibranch suborders, Operation Wallacea survey 1997-1999 (Indonesia). Totally 208 nudibranch species.

Nudibranch genera 1997-99

AeolidinaArmininaDendronotinaDoridina

Figure 5. Proportion of genera within nudibranch suborders, Operation Wallacea survey 1997-1999 (Indonesia). Totally 56 nudibranch genera.

26

Table 2 presents a list of the most common species. Some of these species (and a few others) are illustrated in Plate A and B. Table 2. The most common opisthobranch species during Operation Wallacea survey 1999 (Indonesia). Order Species Observations Occasions Cephalaspidea Chelidonura amoena 24 17 Notaspidea Pleurobranchus forskalii 20 11 Nudibranchia Phidiana indica 17 17 Nudibranchia Phyllodesmium briareum 398 27 Nudibranchia Pteraeolidia ianthina 25 19 Nudibranchia Dermatobranchus sp (OWN 101) 20 13 Nudibranchia Chromodoris annae 215 109 Nudibranchia Chromodoris coi 35 30 Nudibranchia Chromodoris dianae 106 60 Nudibranchia Chromodoris geometrica 49 40 Nudibranchia Chromodoris kuniei 22 17 Nudibranchia Chromodoris leopardus 13 11 Nudibranchia Chromodoris lochi 132 81 Nudibranchia Chromodoris magnifica 119 71 Nudibranchia Chromodoris willani 106 63 Nudibranchia Hypselodoris bullocki 18 13 Nudibranchia Jorunna funebris 29 26 Nudibranchia Nembrotha kubaryana 36 ? Nudibranchia Phyllidia coelestis 113 82 Nudibranchia Phyllidia elegans 164 115 Nudibranchia Phyllidia ocellata 19 19 Nudibranchia Phyllidia varicosa 131 96 Nudibranchia Phyllidiella pustulosa 604 289 Nudibranchia Phyllidiella rudmani 30 24 Nudibranchia Phyllidiopsis pipeki 58 52 Nudibranchia Phyllidiopsis shireenae 12 11 Nudibranchia Phyllidiopsis striata 34 28 Nudibranchia Reticulidia fungia 15 14 Nudibranchia Roboastra gracilis 16 16 Sacoglossa Elysia sp (OWN 99) 15 12 Sacoglossa Thuridilla bayeri 36 25 Sacoglossa Thuridilla hoffae 13 11 Total number 2644

2.4 Discussion Appendix I is the summary of more than 450 hand-written pages with opisthobranch information from the 1999 survey. The material was delivered in three batches between January and April 2000. As numerous people wrote the material, a continuing obstacle was to decipher the notes accurately. Another difficulty was the uses of many synonyms as this made the compilation work very time consuming. A third problem was the fact that the most complete photographic handbook (Debelius, 1996) contains many misidentifications and lacks information about authors.

27

As seen from Figure 1 most of the species (in all orders) were found during 1999. The reason for this is probably because of the increased effort under the 1999 season (May – November). Regrettably, there is no information regarding search effort, which makes it difficult to compare the three years. When looking at the number of genera instead (Figure 2) the picture is less clear. Fewer Cephalaspidean genera were found 1999 than 1997-1998. This is probably because of less search effort during 1999 than 1997-1998. (During my own visit, no actual digging was conducted in search of the burrowing Cephalaspidea.) It is interesting to note that, despite a massive identifying effort, only about 60 % of the opisthobranchs were identified to species level (Figure 3). The majority of the observations involved members of the Nudibranchia, which is not surprising since Nudibranchia is the largest order in Opisthobranchia. As seen from Figures 4 and 5, the most common nudibranch suborder is the Doridina whether looking for species number or generic number. There are very few similar surveys to compare with. The most recent one is the “Nudibranchs of Heron Island, Great Barrier Reef” (Marshall & Willan, 1999). This survey was run between November 1980 and December 1998 and a total of 262 species was found during that period. In comparison 295 species were found during the Operation Wallacea surveys 1997-1999 (Figures 1 and 3). This number will probably be somewhat lower after final examinations but it is still an impressive figure after only three years of surveys.( According to Warren (2000) there are 297 species but I have excluded Colpodaspis thompsoni and Tritonopsilla australis. In the first case because of its uncertain taxonomic status and in the later case because I believe it to be a case of mistaken identity.) The highest count of opisthobranchs for an ongoing survey in the Indo Pacific is 561 species (Madang, Papua New Guinea) but that is for the period 1986 – March 1997 (Gosliner in Marshall & Willan, 1999). If not just looking at surveys then Gosliner & Draheim (1996) give the highest count of Indo Pacific opisthobranchs in a study. They state that there are more than 3400 opisthobranch species in the region and that the locality with the most species, 646, is Papua New Guinea. No doubt, these figures shows that the region of Papua New Guinea and Indonesia is the leading diversity centre for Opisthobranchia. According to Table 2, 75 % of the opisthobranch observations were of the 14 % most common species. One problem is to categorise the found species. Is a species always common when found in large numbers? What if the species is found only a few times but in large number? In my case I choose to regard all species, which were found on more than 10 separate occasions (during the 1999 season) as common. In “Nudibranchs of Heron Island, Great Barrier Reef” (Marshall & Willan, 1999) there is an attempt to give a relative abundance scale. This could function but since the authors do not give information about the time duration of sampling periods it was not possible to use their abundance scale in this report.

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2.5 References Allen, G. R. & Steene, R. 1996. Indo-Pacific coral reef field guide. Tropical Reef Research.

Singapore. Behrens, D. W. 1991. Pacific Coast nudibranchs: a guide to the opisthobranchs, Alaska to

Baja California. Seachallenger. Monterey. Coleman, N. 1989. Nudibranchs of the South Pacific. Neville Coleman’s Sea Australia

Resource Centre. Springwood. Debelius, H. 1996. Nudibranchs and sea snails Indo Pacific field guide. IKAN –

Unterwasserarchiv. Frankfurt. Expedia, 2001. Internet site: www.expedia.com (March 2001). Gosliner, T. 1987. Nudibranchs of Southern Africa. Seachallenger. Monterey. Gosliner, T. M., Behrens, D. W. and Williams, G. C. 1996. Coral Reef Animals of the Indo-

Pacific. Seachallengers. Monterey. Gosliner, T. M. and Draheim, R. 1996. Indo-Pacific opisthobranch gastropod biogeography:

how do we know what we don’t know? American Malacological Bullentin 12: 37-43. Marshall, J. G. & Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef.

Backhuys Publishers. Leiden. Rudman, B. 2000. The Sea Slug Forum. (www.austmus.gov.au/science/invert/mal/forum) Stanzel, K. B. & Newman, H. 1997. Progress report on the 1996 marine survey of the Tukang

Besi (Wakatobi) archipelago, South East Sulawesi, Indonesia. (Internal report: Operation Wallacea).

Warren, L. 2000. Opisthobranchs of the Tukang Besi archipelago, S.E. Sulawesi, 1999 survey report & results. (www.operationwallacea.win-uk.net/opisthob/opisthob.htm (2000-11-15))

Wells, F. E. & Bryce, C. W. 1993. Sea slugs of Western Australia. Western Australian Museum. Perth.

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3 Spatial distribution of nudibranchs according to benthic structure 3.1 Introduction Why do we know so little about nudibranch ecology? One reason is that nudibranch populations are both scarce and transient (Todd, 1981, Marshall & Willan, 1999). This means that finding places where one can do nudibranch research are difficult. One potential site for such research is at Hoga Island (Indonesia) as the nudibranchs are rather abundant at this area. There are also many nudibranch species around Hoga that are quite common and not so shortlived and that makes it possible to do interesting scientific research at this locality. The research facility is run by Operation Wallacea, a joint venture between The Wallacea Development Institute (Indonesia) and Ecosurveys Ltd (United Kingdom). Hoga Island is situated in the Tukang Besi archipelago, South East of Sulawesi. In July 1996 the area was designated as the Wakatobi Marine National Park (Stanzel & Newman, 1997) and since then there has been ongoing research at this locality. My hypothesis was: Nudibranch distribution depends on benthic structure. The reason for the hypothesis is that I wanted to see if it was possible to find environment preferences among some of the more common nudibranchs in the area. 3.2 Material and methods The dive sites, chosen for research, were “Sampela” and “Buoy 3” (Chart 1). Sampela was considered an example of a more polluted area (from a nearby village) and Buoy 3 an example of a more “unspoilt” area. The survey was conducted in 1999 between the eight and twelfth of November (five consecutive days). On both locations we put down two stationary lines, that each were 30 m long. One line was put at “+12 m” and another one at “+3 m”. The “+” sign indicates that the lines were put down on an estimated depth of 3 or 12 m below the reef crest. This was done in accordance with other ongoing projects and relevant literature at hand (English et al. 1994). The actual survey depth varied with the tidal state but we recorded the depth of the starting point on each dive so that we could adjust our measurements later if necessary. On each dive we aimed to spend 20 minutes searching along each line. The search area was roughly 2.5 m above and under the line, thus having a search area of about 150 m2. An alternative search description is to say that we surveyed a volume of 300 m3 (this figure is calculated from the volume of a half cylinder with a radius of 2.5 m). We dived at each line twice a day (before and after lunch), recording observed species, depths and numbers and measuring individual sizes. The number of divers, on each dive, varied between two and five.

30

Chart 1. Location of the dive sites Buoy 3 and Sampela (Tukang Besi, Indonesia) In order to measure the benthic structure, the length of the benthic forms along the line transect was recorded. Standardised codes were used to describe the benthic forms, see Table 1. Collected data were later analysed by using statistical methods such as ANOVA (Analysis of Variance) and DCA (Detrended Correspondence Analysis). Comparable indices regarding benthic structure and diversity of nudibranchs for the two sites were computed by using the Shannon-Weaver Index formula: H = - Σ pi ln pi (Fowler et al., 1998). (The term pi is the proportion of a particular species in a sample which is multiplied by the natural logarithm of itself. H is derived by summing the product for all species in the sample. The minus sign is to make the final value of H positive.)

Buoy 3

Sampela

31

Table 1. Standardised benthic codes. After English et al., 1994. Hard coral Algea Other

ACB = Acropora Branching AA = Algal assembly OT = Other ACE = Acropora Encrusting CA = Coralline algea SC = Soft coral ACD = Acropora Digitate HA = Halimeda SP = Sponge ACS = Acropora Submassive MA = Macro algea SPB = Sponge Branching ACT = Acropora Tabulate TA = Turf algea SPD = Sponge Digitate CB = Coral Branching Abiotic SPE = Sponge Encrusting CD = Coral Digitate DDD = Data missing SPF = Sponge Foliose CE = Coral Encrusting R = Rubble SPM = Sponge Massive CF = Coral Foliose RCK = Rock SPO = Sponge Other form CM = Coral Massive S = Sand SPS = Sponge Submassive CMR = Coral Mushroom SI = Silt SPT = Sponge Tubular, Barrel, CS = Coral Submassive WA = Water Vase DC = Dead coral ZO = Zoanthids DCA = Dead coral With algea CHL = Heliopora CME = Millepora

3.3 Results The results from measuring benthic structure are shown in Table 2. Shannon-Weaver Indices for the four transects were calculated according to Fowler et al., 1998. Table 2. Benthic structure at Buoy 3 and Sampela (Tukang Besi, Indonesia). Benthic code (See Table 1)

Buoy 3 (“+ 3 m”) % Cover

Buoy 3 (“+ 12 m”) % Cover

Sampela (“+ 3 m”) % Cover

Sampela (“+ 12 m”) % Cover

AA 1 2 1 3 ACB 3 CA 7 32 9 20 CB 10 1 CD 10 1 CE 13 15 7 7 CF 2 1 1 CM 1 CMR 3 4 CS 10 DCA 3 3 DC 7 MA 1 OT 2 1 R 12 2 25 11 S 3 38 SC 3 7 13 12 SP 2 5 7 3 SPE 1 4 WA 26 32 3 2 Shannon-Weaver Index

2.251 1.746 2.422 1.780

32

The average line depth and search time (and their standard deviation = SD) for Buoy 3 and Sampela are given in Table 3. Table 3. Average search time and line depth for Buoy 3 and Sampela (Tukang Besi, Indonesia). Buoy 3

AM “+3m”

Buoy 3 PM “+3m”

Buoy 3 AM “+12m”

Buoy 3 PM “+12m”

Sampela AM “+3m”

Sampela PM “+3m”

Sampela AM “+12m”

Sampela PM “+12m)

Average search time (min)

27 27 30 31 32 28 33 31

SD 8 3 11 14 7 6 13 14 Average line depth (m)

4.7 4.7 11.9 14.0 5.5 6.8 12.2 13.3

SD 0.78 0.18 0.22 0.48 0.24 0.05 0.41 0.25 During 40 dives we managed to find a total of 252 nudibranchs belonging to at least 19 species (two nudibranch taxa were not identified to species level). At Sampela we found 138 nudibranchs and at Buoy 3 we found 114. The search record regarding observations and species is presented in Table 4. The complete search record is available in Appendix II. Table 4. Search record for Buoy 3 and Sampela (Tukang Besi, Indonesia). Species Buoy 3 Sampela Chromodoris annae 15 7 Chromodoris boucheti 3 0 Chromodoris coi 0 5 Chromodoris dianae 6 0 Chromodoris elisabethina 5 0 Chromodoris geometrica 1 2 Chromodoris kunei 0 2 Chromodoris leopardus 8 0 Chromodoris magnifica 19 0 Jorunna funebris 17 21 Nembrotha kubaryana 6 0 Reticulidia fungia 3 0 Phyllidiopsis pipeki 5 7 Phyllidiopsis striata 0 9 Phyllidia coelestis 3 15 Phyllidia sp 0 2 Phyllidia varicosa 5 10 Phyllidiella pustolosa 18 55 Unidentified 0 1 Pteraolidia ianthina 0 1 Thecacera picta 0 1

The Shannon-Weaver Index (Fowler et al., 1998) regarding nudibranch species for Buoy 3 is 2.377 and for Sampela 1.976.

33

The effects of the sites (Sampela, Buoy 3), depths (+3m, +12m) and times (“am”, “pm”) were analysed by an ANOVA (By the computer program MANOVA). Both nudibranch observations (Figure 1) and species number (Figure 2) were tested. The lowest p-value (0.08) was found for interaction of sites and depths regarding number of species. In no cases “am/pm” seemed to affect species number or observations.

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34

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Figure 3. Buoy 3 and Sampela (Tukang Besi, Indonesia): Total species number and average (with SE bars) of nudibranchs. Collected data was also used in a Detrended Correspondence Analysis (DCA). This is an indirect gradient analysis technique which can be used for explanatory analysis (Gauch, 1984). The analysis was made with the computer program CANOCO 4.0 (Ter Braak, 1998). The result from this analysis is shown in Figure 4.

35

Figure 4. Results from a “Detrended Correspondence Analysis” (DCA). The data are taken from the sites: Buoy 3 and Sampela (Tukang Besi, Indonesia) 3.4 Discussion Totally, the average number of observed nudibranchs was higher at Sampela (7) than Buoy 3 (6). We found approximately the same average number (6) of nudibranchs around Buoy 3 regardless of depth. At Sampela there was a difference in nudibranch abundance between the

Nembrotha kubyaryana

Reticulidia fungia

Chromodoris leopardus

Chromodoris dianae

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[Cor. encr.,Cor. mushroom ]

36

depths. We found on average more nudibranchs at “+3m” then at “+12m” (Figure 1). The lower number for “+12m” is probably due to the fact that there was a large sandy area at a part of this depth (very few nudibranchs live on sand). At both depths at Buoy 3 the normal pattern was that we would find more chromodorids than phyllidiids. This was not the case at Sampela however, where the situation was reversed. As this reversal occurs whether we look at individual number (Figure 1) or number of species (Figure 2), it would be interesting to check more similar sites and see if there is a larger pattern. There is a possibility that this reversal could work as an environment indicator since we consider Sampela more polluted than Buoy 3. The highest number of species, 14, was found at Buoy 3 but Sampela was not far behind with 12 species. This was in total numbers, when looking at the average values per dive there was no difference between the sites. On each dive we could expect to find about 4 species whether we were diving at Buoy 3 or Sampela (Figure 3). When comparing the “Shannon-Weaver Index” for the two sites (Sampela, 1.976 and Buoy 3, 2.377) there is a cause to say that Buoy 3 seemed to be more diverse than Sampela. As for my hypothesis that “Nudibranch distribution depends on benthic structure”, it is difficult to get a clear picture of what kind of benthic structure the different nudibranchs prefer but according to Figure 4 some of the surveyed nudibranch species are affiliated with specific benthic forms, as Chromodoris dianae, C. annae and C. magnifica occurred with macro algea, digitate and branching coral. Another indication is that Chromodoris dianae, C. annae, C. magnifica, C. coi, C. kunei, Phyllidiella pustolosa, Phyllidia sp and Phyllidiopsis striata seemed to prefer “+3m” to “+ 12m”. Since most nudibranchs are supposed to settle near food sources, I had expected a much more clear and close affiliation between Chromodoridae/Phyllidiidae and the “sponge” categories, as sponges are supposed to be their food source. This connection is not so easy to see in Figure 4. Nevertheless, I still think it should be possible to find out more facts regarding nudibranchs and benthic structure, by doing more survey attempts like the one presented here. 3.5 References English, S., Wilkinson, C. and Baker, V. (eds). 1994. Survey Manual for Tropical Resources.

Australian Institute of Marine Science. Townsville. Fowler, J., Cohen, L. and Jarvis, P. 1998. Practical Statistics for Field Biology. John Wiley

& Sons. Chichester, New York, Weinheim, Brisbane, Singapore, Toronto. Gauch, H. G. Jr. 1984. Multivariate analysis in community ecology. Cambridge University

Press. Cambridge, London, New York, New Rochelle, Melbourne, Sidney. Marshall, J. G. & Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef.

Backhuys Publishers. Leiden. Stanzel, K. B. & Newman, H. 1997. Progress report on the 1996 Marine Survey of the

Tukang Besi (Wakatobi) archipelago, South East Sulawesi, Indonesia. (Internal report: Operation Wallacea).

Ter Braak, C. J. F. 1998. Canoco 4.0 (Computer program). Centre for Biometry Wageningen. Wageningen.

Todd, C. D. 1981. The Ecology of Nudibranch Molluscs. Oceanography and Marine Biology: an Annual Review 19: 141-234.

37

4 Conclusions and recommendations In section 3, I had another hypothesis: Chromodorids have larger home ranges than Phyllidids. The background for this hypothesis was that there was a general field observation among the volunteers that nudibranchs of the family Phyllidiidae were much more stationary than nudibranchs of the family Chromodorididae. Accordingly, I wanted to objectively test this observation. We first tried to use a coordinate system and write down the co-ordinates for every nudibranch we found, hoping to be able to follow their movement. It seemed to work at first but later a storm ruined our efforts by moving the lines a little bit and thus made it impossible to continue using co-ordinates. We then tried to mark the nudibranch position with a string of rope and a buoyant bamboo marker. This also seemed to work at first but we later got problem as the bamboo markers lost their buoyancy and that made it very difficult to find them. It was also difficult to recognise the specific nudibranchs and I believe that we would have needed an identification system in order to successfully follow the nudibranchs. Perhaps a “fish marking” system with tags would work? Another problem is to keep a functioning “dive relay race” going on for a long enough period. When doing my own survey we also discovered how difficult it is to do scientific work underwater. As we had to be several divers sharing the workload we always had to adjust for the least experienced and/or the diver with the highest air consumption. This makes underwater surveys with volunteers extremely difficult due to the differences in diving experience, biological knowledge and air consumption. It takes many dives before you can find a system that is workable. Another problem with working in a remote area like Wakatobi is the lack of equipment and the logistic transport problems when needing repairs or spare parts. This is especially hard since the climate affects and wears down all kinds of equipment, for instance, diving equipment, boats, recording devices and of course computer equipment. There is always the need to come up with new repair ideas, using very simple tools or equipment. We also found out how hard the tropical environment affects the personnel, usually about one third are on the sick list due to infections (stomach illness, ear infections, badly infected wounds, etc). This means that you always have to have a lot more people helping you than you would normally expect. In the future I think it would be worthwhile to do a more structured opisthobranch study at Hoga. One main mistake, when working with the checklist (section 2), is that there has been no registration of search effort. This makes it impossible to correlate the opisthobranch abundance with, for instance, dive time or search area. The easiest way to register the search effort is to keep track of the “active search time” but another idea could be to cooperate with other projects where the search area is defined by transects. In these cases an estimation of search area is given for “free”!

38

5 Acknowledgements I would like thank: Per Milberg, for all the help with this report. - I could not have done it without your help. Lindsay Warren and Krzystof Brzkieta, for trying to help out at Hoga when my appointed (Opwall) supervisor disappeared! All the volunteers at Hoga who graciously tried to help me, especially my dive buddies: Peter Bächi, Rob Saunders, Nikki Rowland, Jenny Hill, Elaine Worley, Gail Young, Sarah Yeates and Tony Gatcombe. Richard Willan, Pam Beesley, The Ray Society, David Behrens and the Linnean Society of London for kind permissions to reproduce appropriate illustrations in my thesis. My parents, for financial and moral support. Åsa Nilsson, my fiancé, for helping me, all the time. – Thank You!

Plate A

Chelidonura amoena Bergh, 1905 Pleurobranchus grandis Pease, 1868 Note: White squid cleaning the opisthobranch!

Cyerce elegans Bergh, 1870 Nembrotha sp (lineolata? Bergh, 1905)

Jorunna funebris (Keelart, 1858) Hexabranchus sanguineus (Rüppell & Leuckart, 1828) Note: The identification details are taken from “The Sea Slug Forum” (www.seaslugforum.net 2001-04-22). Copyright holder and photographer is the author (Lars Karlsson). The photos are all from Hoga Island (1999).

Plate B

Chromodoris annae Bergh, 1877 Chromodoris magnifica (Quoy & Gaimard, 1832)

Chromodoris sp (boucheti? Rudman, 1982) Phyllidiella pustulosa (Cuvier, 1804)

Phyllidia sp (willani? Brunckhorst, 1993) Phyllidiopsis striata Bergh, 1888 Note: The identification details are taken from “The Sea Slug Forum” (www.seaslugforum.net 2001-04-22). Copyright holder and photographer is the author (Lars Karlsson). The photos are all from Hoga Island (1999).

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929)

11

9906

0914

8&

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N 1

48F

labe

llini

dae

Fla

belli

nasp

. 1.

x1

198

1112

9906

0567

Aeo

lidiid

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men

andr

ano

dosa

Hae

felfi

nger

& S

tam

m, 1

958

11

9811

2499

0906

77G

lauc

idae

Fac

elin

idae

Mor

idill

abr

ocki

Ber

gh, 1

888

9803

256

Gla

ucid

aeF

acel

inid

aeP

hidi

ana

indi

ca(B

ergh

, 189

6)17

1698

1019

9905

1599

1116

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alor

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Gla

ucid

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acel

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desm

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bria

reum

(Ber

gh, 1

896)

398

2798

0916

9905

1799

1121

(bria

reus

)G

lauc

idae

Fac

elin

idae

Phy

llode

smiu

mlo

ngic

irrum

(Ber

gh, 1

905)

11

9709

2099

0721

Phy

llode

smiu

msp

33

9906

2399

0816

124

sim

ilar

to O

WN

85

Phy

llode

smiu

msp

9709

2085

Aeo

lidiid

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atra

Bab

a, 1

955

9811

0465

Gla

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aeF

acel

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nthi

na(A

ngas

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4)25

1997

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9905

0599

1111

Aeo

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1202

82A

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9812

0283

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lid1

199

0617

117

Aeo

lid1

199

0630

126

Aeo

lid2

199

0630

127

Aeo

lid1

199

0617

129

Aeo

lid1

199

0816

174

Su

bo

rder

Aeo

lidin

aA

rmin

idae

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mat

obra

nchu

scf

. gon

atop

hora

x97

0920

D. g

onat

opho

ra in

"D

" 19

96 p

295

.D

erm

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ranc

hus

sp3

398

0907

9905

1599

0917

51D

erm

atob

ranc

hus

sp20

1399

0603

9906

1510

1D

erm

atob

ranc

hus

sp2

199

1111

206

Der

mat

obra

nchu

ssp

9811

1368

Der

mat

obra

nchu

ssp

11

9905

1490

Der

mat

obra

nchu

ssp

11

9906

2111

9D

erm

atob

ranc

hus

sp1

199

0805

165

Der

mat

obra

nchu

ssp

11

9910

2219

8D

erm

atob

ranc

hus

sp1

199

0624

cf O

WN

101

A

ppen

dix

I, 3(

6)

Com

pila

tion

over

Opi

stho

bran

chs

foun

d du

ring

Ope

ratio

n W

alla

cea

surv

ey 1

997-

1999

.A

rmin

idae

Der

mat

obra

nchu

ssp

. 2.

x98

1124

74S

ub

ord

er A

rmin

ina

Bor

nelli

dae

Bor

nelli

dae

Bor

nella

angu

illa

John

son,

198

32

297

0920

9905

1599

0517

Bor

nelli

dae

Bor

nelli

dae

Bor

nella

stel

lifer

(Ada

ms

& R

eeve

, 184

8)33

599

1010

9911

24In

"G

BR

O":

(A

. Ada

ms

& R

eeve

in A

. Ada

ms,

184

8)D

endr

onot

idae

Den

dron

otus

spx

33

9709

2099

0610

9908

2516

8T

riton

idae

Mar

ioni

asp

.x

9803

2513

1O

WN

13/1

Mar

ioni

avi

rides

cens

22

9805

0199

0607

9907

03T

ethy

dida

eM

elib

een

geli

Ris

bec,

193

71

199

1102

203

Tet

hydi

dae

Mel

ibe

fimbr

iata

Ald

er &

Han

cock

, 186

43

299

0903

9909

15S

cylla

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eS

cylla

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eS

cylla

eape

lagi

ca(L

inne

aeus

, 175

8)1

199

0730

In "

GB

RO

": L

inne

, 175

8Tr

itoni

asp

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0325

12Tr

itoni

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199

0708

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alba

(Bab

a, 1

949)

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silla

)(T

riton

opsi

llaau

stra

lis)

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roba

bly

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isst

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bo

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Den

dro

no

tin

aC

hrom

odor

idid

aeC

hrom

odor

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aeA

rdea

doris

egre

ttaR

udm

an, 1

984

87

9805

0199

0603

9911

20D

orid

idae

Ast

eron

otus

cesp

itosu

s(H

asse

lt, 1

824)

11

9709

2099

0621

Chr

omod

orid

idae

Cer

atos

oma

mia

mira

na(B

ergh

, 187

5)2

299

0717

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hrom

odor

idid

aeC

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lt, 1

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11

9911

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hrom

odor

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nue

Abr

aham

, 187

61

199

0824

Chr

omod

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omod

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atos

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trilo

batu

m(J

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ray,

182

7)98

0901

In "

GB

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Gra

y, 1

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Chr

omod

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omod

orid

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Chr

omod

oris

albo

punc

tata

(Gar

ret,

1879

)1

199

0731

157

Chr

omod

oris

aliu

s1

199

0914

In "

WB

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ecie

s 14

7. R

udm

an 1

987

Chr

omod

orid

idae

Chr

omod

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anna

eB

ergh

, 187

721

510

997

0920

9905

0599

1117

Chr

omod

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idae

Chr

omod

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bouc

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Rud

man

, 198

28

698

0501

9907

0399

1001

(bou

chet

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odor

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i(R

isbe

c, 1

956)

3530

9709

2099

0524

9911

05C

hrom

odor

idid

aeC

hrom

odor

isdi

anae

Gos

liner

& B

ehre

ns, 1

998

106

6097

0920

9905

1499

1117

Chr

omod

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idae

Chr

omod

orid

idae

Chr

omod

oris

elis

abet

hina

Ber

gh, 1

877

98

9709

2099

0524

9909

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lizab

ethi

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Chr

omod

orid

idae

Chr

omod

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idae

Chr

omod

oris

fidel

is(K

elaa

rt, 1

858)

33

9905

2199

0722

Chr

omod

orid

idae

Chr

omod

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idae

Chr

omod

oris

geom

etric

aR

isbe

c, 1

928

4940

9709

2099

0603

9911

20C

hrom

odor

idid

aeC

hrom

odor

idid

aeC

hrom

odor

ishi

ntua

nens

is97

0920

Chr

omod

orid

idae

Chr

omod

orid

idae

Chr

omod

oris

kuite

riR

udm

an, 1

982

11

9905

22ku

iteri

? O

S 0

184

Chr

omod

orid

idae

Chr

omod

orid

idae

Chr

omod

oris

kuni

eiP

ruvo

t-F

ol, 1

930

2217

9709

2099

0518

9911

09C

hrom

odor

idid

aeC

hrom

odor

idid

aeC

hrom

odor

isle

opar

dus

Rud

man

, 198

713

1197

0920

9906

0999

1104

Chr

omod

orid

idae

Chr

omod

orid

idae

Chr

omod

oris

loch

iR

udm

an, 1

982

132

8197

0920

9905

0599

1117

Chr

omod

oris

cf. l

ochi

11

9906

2312

3C

hrom

odor

idid

aeC

hrom

odor

idid

aeC

hrom

odor

ism

agni

fica

(Quo

y &

Gai

mar

d, 1

832)

119

7197

0920

9905

2099

1117

Chr

omod

orid

idae

Chr

omod

oris

prec

iosa

(Kel

aart

, 185

8)1

199

0704

Chr

omod

oris

quad

ricol

our

9808

01In

Deb

eliu

s 19

96 p

200

Chr

omod

oris

retic

ulat

a2

298

0315

9911

0199

1111

In "

GB

W"

spec

ies

577

(Quo

y &

Gai

mar

d, 1

832)

(=t

inct

oria

)C

hrom

odor

issp

32

9907

1199

1007

142

Chr

omod

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sp98

1126

81C

hrom

odor

issp

11

9906

0310

0C

hrom

odor

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11

9907

1514

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hrom

odor

issp

11

9907

2414

9C

hrom

odor

idid

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hrom

odor

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aeC

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odor

isst

rigat

aR

udm

an, 1

982

54

9803

1899

0802

9911

172

Chr

omod

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idae

Chr

omod

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verr

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(Cro

sse,

187

5)2

298

0831

9907

0499

0730

48ve

rrie

ri? a

lbon

ares

Chr

omod

orid

idae

Chr

omod

oris

will

ani

Rud

man

, 198

210

663

9804

0199

0505

9911

16D

endr

odor

idid

aeD

endr

odor

isca

rbun

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sa(Q

ouy

& G

aim

ard,

183

2)1

199

0906

179

Den

drod

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idae

Den

dodo

ridid

aeD

endr

odor

isni

gra

(Stim

pson

, 185

5)98

0801

Den

drod

oris

cf. n

igra

11

9910

1819

6D

endr

odor

idid

aeD

endo

dorid

idae

Den

drod

oris

tube

rcul

osa

(Quo

y &

Gai

mar

d, 1

832)

9804

04(D

endr

ois)

Den

drod

oris

?98

1216

86D

endr

odor

is ?

11

9911

0120

1D

iaph

orod

oris

mits

ui(B

aba,

193

8)98

1119

69In

Gos

liner

,198

7 (f

orm

er D

. Sp)

Dor

idid

aeD

isco

doris

boho

liens

isB

ergh

, 187

71

199

0728

A

ppen

dix

I, 4(

6)

Com

pila

tion

over

Opi

stho

bran

chs

foun

d du

ring

Ope

ratio

n W

alla

cea

surv

ey 1

997-

1999

.D

orid

idae

Dis

codo

ris?

litur

ata

Ber

gh, 1

905

21

9808

1599

0721

45(D

isco

doris

? li

tura

ta)

Dor

idid

aeD

oris

?sp

. 10.

x2

299

0702

9907

1713

1D

oris

sp.

10

Chr

omod

orid

idae

Dur

ville

doris

pusi

llax

11

9907

1814

7C

hrom

odor

idid

aeD

urvi

lledo

rissi

mila

risx

11

9908

1116

9Fr

yeria

men

indi

eB

runc

khor

st, 1

993

87

9709

2099

0603

9909

01(m

eini

ndae

)Fr

yeria

ruep

pelii

Ber

gh, 1

869

11

9906

21(c

f Fry

eria

rup

peli)

Glo

ssod

oris

albo

cinc

ta98

0620

Chr

omod

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omod

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ssod

oris

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ta(B

ergh

, 188

8)98

0620

Chr

omod

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idae

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omod

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idae

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ssod

oris

hiku

eren

sis

(Pru

vot-

Fol

, 195

4)2

298

1024

9906

0599

0806

Chr

omod

orid

idae

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omod

orid

idae

Glo

ssod

oris

rufo

mar

gina

ta(B

ergh

, 189

0)98

1216

Gym

nodo

ridid

aeG

ymno

doris

alba

(Ber

gh, 1

877)

11

9909

11G

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dorid

idae

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nodo

ridid

aeG

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doris

ceyl

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a(K

elaa

rt, 1

858)

53

9911

0299

1105

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auth

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sp.

11

9911

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8G

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dorid

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Gym

nodo

rissp

. 1.

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1016

58D

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Dor

idid

aeH

alge

rda

carls

oni /

aff.

car

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iR

udm

an, 1

978

11

9709

2099

1111

carls

oni

Dor

idid

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alge

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mal

esso

Car

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& H

off,

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22

9806

2099

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9910

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in D

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1996

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104

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9709

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ll &

Leu

ckar

t, 18

30)

55

9810

0199

0527

9911

1619

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(R

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ll &

Leu

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omod

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omod

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selo

doris

bullo

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(Col

lingw

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188

1)18

1397

0920

9905

2499

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smal

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omod

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omod

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selo

doris

emm

a / e

mm

aeR

udm

an, 1

977

9709

20em

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omod

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selo

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infu

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(Rup

pel &

Leu

ckar

t, 18

28)

11

9810

1699

0825

57H

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lodo

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nsso

n, 1

999

11

9907

2815

4In

ZJL

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Chr

omod

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omod

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idae

Hyp

selo

doris

mac

ulos

a(P

ease

, 187

1)5

599

0520

9909

15C

hrom

odor

idid

aeH

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lodo

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ulos

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11

9907

0713

4C

hrom

odor

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lodo

rispu

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Ham

atan

i, 19

951

198

0421

9908

0920

Chr

omod

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selo

doris

rudm

ani

x1

199

0910

183

Hyp

selo

doris

sp.

11

9908

1117

0C

hrom

odor

idid

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. 1.

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1103

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dam

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11

9908

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3D

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idid

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runn

afu

nebr

is(K

elaa

rt, 1

858)

2926

9709

2099

0513

9911

18Jo

runn

asp

.1

199

1026

199

Pol

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brot

hach

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rlain

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Beh

rens

, 199

797

0920

Pol

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Pol

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Nem

brot

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ergh

, 187

710

9709

2099

0525

9910

26P

olyc

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kuba

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, 187

736

9709

2099

0524

9911

11P

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gh, 1

905

497

0920

9910

1599

1117

Pol

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idae

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brot

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tilan

s(P

ruvo

t-F

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931)

297

0920

9911

0999

1116

Nem

brot

hasp

. 97

0920

Pol

ycer

idae

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brot

hasp

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9811

2476

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iritid

aeA

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dae

Not

odor

isga

rdin

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Elio

t, 19

0398

0915

Aeg

iritid

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Not

odor

ism

inor

Elio

t, 19

0498

0324

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odor

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rena

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0606

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spec

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610.

(G

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Beh

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199

8)C

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man

, 198

697

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Chr

omod

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idae

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mea

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(Pea

se, 1

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11

9908

1617

1va

rians

/ no

rba

Chr

omod

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mea

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299

0630

9907

3012

8G

onio

dorid

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nia

sp.

x97

0920

(=H

opki

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)P

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nodo

risau

rora

9810

2964

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e M

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odor

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odor

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nodo

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ta(A

. Ada

ms

& R

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185

0)2

298

0706

9906

0699

0806

103

& O

WN

103,

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llidi

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llidi

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llidi

aba

bai

Bru

nckh

orst

, 199

33

297

0920

9907

2799

0802

Phy

llidi

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llidi

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Phy

llidi

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, 190

511

382

9709

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0514

9911

17

A

ppen

dix

I, 5(

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Com

pila

tion

over

Opi

stho

bran

chs

foun

d du

ring

Ope

ratio

n W

alla

cea

surv

ey 1

997-

1999

.P

hylli

dia

sp. c

f. co

eles

tis1

199

0807

167

Phy

llidi

idae

Phy

llidi

idae

Phy

llidi

ael

egan

sB

ergh

, 186

916

411

597

0920

9905

1999

1117

Phy

llidi

idae

Phy

llidi

idae

Phy

llidi

aex

quis

itaB

runc

khor

st, 1

993

98

9709

2099

0721

9911

05P

hylli

diid

aeP

hylli

diid

aeP

hylli

dia

ocel

lata

Cuv

ier,

180

419

1997

0920

9906

2499

1105

Phy

llidi

asp

33

9907

0999

0812

140

Phy

llidi

asp

11

9906

2313

0P

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dia

sp1

199

0708

136

Phy

llidi

asp

66

9907

1299

1115

138

9907

08 +

1 ?

Phy

llidi

asp

11

9907

0913

9P

hylli

dia

sp1

199

0713

143

Phy

llidi

asp

11

9907

1414

4P

hylli

dia

sp1

199

0729

153

Phy

llidi

idae

Phy

llidi

atu

laM

arcu

s &

Mar

cus,

197

01

199

0707

Phy

llidi

idae

Phy

llidi

idae

Phy

llidi

ava

ricos

aLa

mar

ck, 1

801

131

9697

0920

9905

1599

1117

Phy

llidi

idae

Phy

llidi

aw

illan

iB

runc

khor

st, 1

993

76

9810

1799

0628

9911

0959

Phy

llidi

idae

Phy

llidi

idae

Phy

llidi

ella

nigr

a(v

an H

asse

lt, 1

824)

154

9809

0199

0515

9909

17In

"G

BR

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(H

asse

lt, 1

824)

. Lah

oa C

hann

elP

hylli

diid

aeP

hylli

diid

aeP

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diel

lapu

stul

osa

(Cuv

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180

4)60

428

997

0920

9905

0599

1124

Phy

llidi

idae

Phy

llidi

idae

Phy

llidi

ella

rudm

ani

Bru

nckh

orst

, 199

330

2498

1002

9906

1999

1028

Phy

llidi

idae

Phy

llidi

ella

zeyl

anic

a(K

elaa

rt, 1

859)

108

9906

2299

1112

Phy

llidi

idae

Phy

llidi

opsi

san

nae

Bru

nckh

orst

, 199

32

299

0802

9908

05P

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diid

aeP

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diop

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daut

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(Vay

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re, 1

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11

9910

1819

5P

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diop

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fissu

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runc

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st, 1

993

11

9909

3019

3P

hylli

diid

aeP

hylli

diid

aeP

hylli

diop

sis

krem

pfi

Pru

vot-

Fol

, 195

72

299

0917

9911

0118

5P

hylli

diid

aeP

hylli

diid

aeP

hylli

diop

sis

pipe

kiB

runc

khor

st, 1

993

5852

9709

2099

0519

9911

24P

hylli

diid

aeP

hylli

diid

aeP

hylli

diop

sis

shire

enae

Bru

nckh

orst

, 199

312

1197

0920

9906

2699

1113

(shi

rinae

)P

hylli

diid

aeP

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diop

sis

sphi

ngis

Bru

nckh

orst

, 199

31

199

0828

Phy

llidi

idae

Phy

llidi

idae

Phy

llidi

opsi

sst

riata

Ber

gh, 1

888

3428

9709

2099

0604

9911

18D

orid

idae

Pla

tydo

risfo

rmos

a(A

lder

& H

anco

ck, 1

864)

52

9905

2099

1103

Pla

tydo

rissc

abra

(Cuv

ier,

180

4)7

499

0716

9910

27P

laty

doris

sp2

199

0820

190

Pol

ycer

idae

Trio

phid

aeP

loca

mop

heru

sce

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ica

/ cey

loni

cus

(Kel

aart

, 185

8)3

299

0906

9909

2518

1ce

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Pol

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idae

Pol

ycer

asp

. 3.

x1

199

0920

188

Pol

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a ?

9812

0284

Phy

llidi

idae

Phy

llidi

idae

Ret

icul

idia

fung

iaB

runc

khor

st &

Gos

liner

, 199

315

1497

0920

9906

1999

1118

In "

GB

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": B

runc

khor

st &

Gos

liner

in B

runc

khor

st, 1

993

Phy

llidi

idae

Phy

llidi

idae

Ret

icul

idia

halg

erda

Bru

nckh

orst

& B

urn,

199

01

197

0920

9906

09In

"G

BR

O":

Bru

nckh

orst

& G

oslin

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Bru

nckh

orst

, 199

3C

hrom

odor

idid

aeR

isbe

cia

impe

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(Pea

se, 1

860)

9809

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hrom

odor

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hrom

odor

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aeR

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cia

tryo

ni(G

arre

tt, 1

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97

9803

1999

0609

9911

13P

olyc

erid

aeR

oboa

stra

grac

ilis

(Ber

gh, 1

877)

1616

9709

2099

0515

9911

12D

orid

idae

Seb

ador

isnu

bilo

saP

ease

, 189

11

199

0521

94P

olyc

erid

aeP

olyc

erid

aeT

ambj

am

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a(B

ergh

, 187

7)97

0920

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9709

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(Ber

gh, 1

883)

9709

20P

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aeP

olyc

erid

aeT

heca

cera

pict

aB

aba,

197

22

297

0920

9910

2299

1111

Pol

ycer

idae

The

cace

rasp

x97

0920

Chr

omod

orid

idae

Tho

runn

aau

stra

lis(R

isbe

c, 1

928)

9811

21C

hrom

odor

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aeC

hrom

odor

idid

aeT

horu

nna

furti

vaB

ergh

, 187

813

898

1119

9906

0399

1116

71T

rapa

nia

sp.

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2630

Gon

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rapa

nia

sp. 2

.x

9811

2475

Gon

iodo

ridid

aeG

onod

orid

idae

Tra

pani

ato

ddi

Rud

man

, 198

71

199

0617

Tra

pani

a ?

11

9907

0413

2T

rapa

nia

?1

199

0728

151

Dor

idid

aeD

orid

idae

Trip

pa

inte

cta

(Kel

aart

, 185

8b)

22

9809

0999

0613

9911

16T

rippa

sp

32

9906

1499

0903

112

Trip

pa

sp98

0530

34C

hrom

odor

id98

0624

39C

hrom

odor

id1

199

0515

91

A

ppen

dix

I, 6(

6)

Com

pila

tion

over

Opi

stho

bran

chs

foun

d du

ring

Ope

ratio

n W

alla

cea

surv

ey 1

997-

1999

.S

ub

ord

er D

ori

din

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ia28

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rmin

ina

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, Den

dro

no

tin

a =

47, D

ori

din

a =

2328

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Cos

tasi

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sp1

198

0621

9906

2112

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osta

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lidae

Cos

tasi

ella

sp. 1

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9811

2480

Cos

tasi

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aeC

osta

siel

la ?

sp. 2

x1

199

0518

93C

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hylli

dae

Pol

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nchi

idae

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rce

eleg

ans

Ber

gh, 1

870

11

9911

1720

7In

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BR

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gh, 1

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54

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0617

9909

10E

lysi

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sia

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x98

1019

62E

lysi

asp

11

9810

2999

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63E

lysi

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9905

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lysi

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32

9906

499

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105

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299

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11

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aeE

lysi

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180

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kobr

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lapu

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Ber

gh, 1

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9906

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Pla

kobr

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9909

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1599

0515

9911

15In

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kobr

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9809

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2199

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9909

14In

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hurid

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0517

9906

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9906

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11

9811

1699

0805

In "

GB

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spec

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550.

Gos

liner

199

5O

rder

= S

aco

glo

ssa

159

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Fam

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natio

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cord

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(in M

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386

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5 43

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(min

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dep

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9911

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Dep

th (

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H-C

(cm

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cm)

Su

rfac

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olo

ur

Sp

ecie

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(mm

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epth

(m

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cm)

V-C

(cm

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urf

ace

Co

lou

rC

hrom

odor

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lisab

ethi

na25

6,6

300

290

CA

Chr

omod

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ann

ae40

719

064

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SO

rang

eC

hrom

odor

is m

agni

fica

506,

143

023

0D

CA

Chr

omod

oris

ann

ae36

719

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SP

SO

rang

eJo

runn

a fu

nebr

is20

4,6

1480

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SP

LP

hylli

diel

la p

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205

680

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PE

Gre

yJo

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nebr

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4,6

1480

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SP

LP

hylli

diel

la p

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205

680

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PE

Gre

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Div

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rch

time

(min

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ine

dep

th (

m)

Dat

eD

iver

sS

earc

htim

e (m

in)

Lin

e d

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(m

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4,7

9911

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pec

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Dep

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H-C

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rfac

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(mm

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(m

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Co

lou

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(cm

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kub

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5,8

1080

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cuc

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odor

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256,

419

05

AA

0Jo

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nebr

is35

5,2

1490

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SP

Chr

omod

oris

ann

ae25

6,4

190

5A

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Joru

nna

fune

bris

205,

214

90-5

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PC

hrom

odor

is m

agni

fica

607,

220

01

SP

0C

hrom

odor

is e

lisab

ethi

na28

6,1

1640

56C

EB

row

nC

hrom

odor

is m

agni

fica

404,

242

010

She

ll5

NP

hylli

dia

varic

osa

354,

824

60-5

6C

AP

ink

Joru

nna

fune

bris

305,

114

8511

SP

9 N

Joru

nna

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bris

205,

114

8511

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9 N

Dat

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Lin

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Div

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time

(min

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dep

th (

m)

9911

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H/S

Y/E

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9911

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H/S

Y/E

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(mm

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Mar

ker

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olo

ur

Dis

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cm)

Sp

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(mm

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Chr

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18 N

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354,

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hylli

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la p

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255,

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29 S

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hrom

odor

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35 S

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11C

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row

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Phy

llidi

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pus

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7 S

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EB

row

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nna

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305,

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8511

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WP

hylli

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306,

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5023

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11C

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row

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llidi

a va

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5,5

2250

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8 N

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Div

ers

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time

(min

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dep

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earc

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9911

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m)

Dep

th (

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(cm

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pec

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e (m

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th (

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(cm

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lou

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(cm

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odor

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407,

16

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omod

oris

mag

nific

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520

032

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10 N

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mod

oris

elis

abet

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304,

911

017

MA

70 N

EP

hylli

diel

la p

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595

012

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NE

Chr

omod

oris

mag

nific

a30

3,1

380

8037

NW

Joru

nna

fune

bris

4520

602

CE

0C

hrom

odor

is m

agni

fica

453,

867

069

60 N

WP

hylli

dia

varic

osa

2250

239

NW

Phy

llidi

a co

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tis30

523

0023

DC

8 E

Phy

llidi

ella

pus

tulo

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6,4

2980

3610

SW

Dat

eD

iver

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earc

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Lin

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time

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dep

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