ant-like stone beetles, ants, and their associations (coleoptera: scydmaenidae; hymenoptera:...

Ant-Like Stone Beetles, Ants, and Their Associations (Coleoptera: Scydmaenidae; Hymenoptera:Formicidae; Isoptera)Author(s): Sean T. O'KeefeSource: Journal of the New York Entomological Society, Vol. 108, No. 3/4 (Fall - Winter,2000), pp. 273-303Published by: New York Entomological SocietyStable URL: http://www.jstor.org/stable/25010360 .

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J. New York Entomol. Soc. 108(3-4):273-303, 2000

ANT-LIKE STONE BEETLES, ANTS, AND THEIR ASSOCIATIONS (COLEOPTERA: SCYDMAENIDAE;

HYMENOPTERA: FORMICIDAE; ISOPTERA)

SEAN T. O'KEEFE

Department of Entomology, Texas A&M University, College Station, Texas 77843

Abstract.-The literature on ant-like stone beetle-ant associations is reviewed and tabulated,

bringing together nearly all published accounts of associations between these two groups. One hundred-seventeen species of Scydmaenidae (Coleoptera) in 20 genera are associated with 45

species of ants (Hymenoptera: Formicidae) in 28 genera. In addition, 16 species of Scydmaen

idae are known associates of termites (Isoptera).

Key words: Scydmaenidae, Formicidae, myrmecophiles, ants, ant-associates, termitophiles.

Associates of ants are numerous and diverse and have a long history of study (Holldobler and Wilson, 1990). Wasmann (1894) published the first comprehensive review of myrmecophiles and developed the first system of categorizing them based on behavior. His classification divided myrmecophiles into five behavioral categories: synechthrans (treated offensively by their hosts), synoeketes (primarily scavengers and predators, ignored by hosts), symphiles (accepted by hosts as guests), ectopar asites and endoparasites (parasites on hosts), and trophobionts (supply hosts with food, e.g., aphids, scale insects, etc.). Donisthorpe, in his book on British myrme cophilous insects (Donisthorpe, 1927b), grouped the first four behaviors proposed by Wasmann into passive relationships and placed trophobionts as active relation ships. Franc, in his work on Slovakian myrmecophilous beetles (Franc, 1992), di vided myrmecophilic behavior into three categories: synecthricans, synoecious myr

mecophiles (synoeketes of Wasmann), and symphilic myrmecophiles (symphiles of

Wasmann). Wilson (1971), Kistner (1982), and Holldobler and Wilson (1990) give recent and excellent reviews on the general biology of myrmecophiles.

Numerous arthropods have been associated with ants. Among non-insect arthro

pods, isopods, millipedes, pseudoscorpions, and numerous species of spiders and mites are known to be myrmecophiles and among the insects, myrmecophiles include species of Collembola, Thysanura, Orthoptera, Blattodea, Psocoptera, Hemiptera,

Homoptera, Neuroptera, Lepidoptera, Diptera, Strepsiptera, Hymenoptera, and Co

leoptera (Wilson, 1971; Kistner, 1982; Holldobler and Wilson, 1990). In the Cole

optera, myrmecophily is widely represented. Kistner (1982) listed 37 families and

Holidobler and Wilson (1990) listed 35 families of beetles associated with ants. Beetle myrmecophily is relatively common (found in at least 15 genera) in Carabi

dae, Histeridae, Leiodidae, Scarabaeidae, Staphylinidae, and Tenebrionidae and fairly prevalent (found in at least four or five genera) in Chrysomelidae, Colydiidae, Cur

culionidae, Limulodidae, Ptiliidae, and Ptinidae (Hoildobler and Wilson, 1990). Sev eral myrmecophilous species in Pella, Dinarda, Tetradonia, Ecitomorpha, Ecito phya, Atemelus, and Limechusa (Staphylinidae: Aleocharinae), Claviger and Adranes

(Staphylinidae: Pselaphinae), Hetaerius (Histeridae), and Cremastocheilus and Myr



274 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 108(3-4)

mecaphodius (Scarabaeidae) are relatively well studied. Beetle-ant associations have

been published for Australia (Lea, 1910, 1912), England (Donisthorpe, 1927b), west

ern Europe (Bernard, 1968), and Slovakia (Franc, 1992; Hlavac and Lackner, 1998). Far more detailed information on the diversity of beetle myrmecophiles is found in

Kistner's compendium (Kistner, 1982) and an excellent review of some of the well

studied myrmecophile biology is found in Holldobler and Wilson's 1990 text.

Although numerous scydmaenid-ant associations have been reported in the liter ature (Table 1), they are widely scattered and usually embedded in more obscure taxonomic works; none were listed by Holldobler and Wilson (1990). Wasmann

(1894) provided the only world-wide review of scydmaenid-ant associations, but it

is considerably out of date. More recently, Peez (1957) listed the scydmaenid-ant

associations known for Italy, Donisthorpe (1927b) included short discussions for those scydmaenids known to be associates of ants in England, and Franc (1992) and

Hlavack and Lackner (1998) listed those known for Slovakia. It is my hope that by

gathering the obscurely published accounts of scydmaenid-ant associations here that others who work more intimately with ants will take a closer look at these poorly known associates.

MATERIALS AND METHODS

All literature at hand on scydmaenid beetles (some 1,000 references) was scanned for mention of associations with ants and entered into a spreadsheet to generate Tables 1 and 2. Names of ants were then updated according to Bolton (1995), and

scydmaenid generic names were updated according to Newton and Franz (1998). Also, notes were taken of any accounts where scydmaenid beetles were associated with termites.

RESULTS

Sixty-six references were found that mentioned scydmaenid-ant associations. Sev

eral of these (e.g., Bernard, 1968; Everts, 1903; Porta, 1957; Reitter, 1909) merely

repeat previously published accounts, but most either mention the association with

ants as part of original descriptions of species or are smaller studies looking specif

ically at myrmecophilous beetles. Twenty-eight genera and 45 species of ants were

associated with 20 genera and 117 species of scydmaenid beetles (Tables 1, 2). Sixteen species of scydmaenids were associated with termites (Table 3), and, unfor

tunately, in nearly every case the species of termite was not identified.

DISCUSSION

Scydmaenid biology. The biology of scydmaenid beetles is still poorly known.

These beetles usually occur in moist forest leaf litter or are sometimes found under

rocks or under bark. In several genera (e.g., Lophioderus Casey, Veraphis Casey,

Brachycepsis Brendel, etc.) the females are either brachypterous or apterous; in other

genera (e.g., Papusus Casey) both males and females are apterous. Most scydmaen ids that I have seen collected in flight intercept traps are males, which suggests that

males of many scydmaenid species are the more active dispersers. Scydmaenids are

infrequently encountered, and they are usually not abundant, although I have found



2000 ASSOCIATIONS OF ANT-LIKE STONE BEETLES (COLEOPTERA) 275

literally hundreds of individuals of a species of Scydmaenus Latreille congregated

under leaves near streams in Guanacaste, Costa Rica during the dry season. Schmid (1988) published the only comprehensive study about scydmaenid biol

ogy, in which he described their prey and feeding strategies. He found that the majority of scydmaenid beetles he studied (about 20 European species) are specialist predators on oribatid mites (Schmid, 1988). He described two methods of prey cap ture and discussed the modifications of the mandibles for these specialized tech niques. The typical scydmaenid mandible has a single, large incisor and often no

subapical teeth (see O'Keefe, 1997a, b, c; Schmid, 1988 for examples). Schmid (1988) described how the mandibles are modified and used like a can opener to pry open either the genital or anal plates of mites. However, members of the Clidicini

(Clidicus Laporte and Papusus) and possibly also members of Scydmaenus are either generalist predators or specialize on prey other than mites because the mandibles of

most species have at least two or three large subapical teeth. Members of one species of Clidicus were found feeding on collembolans (O'Keefe and Monteith, in prepa ration) and members of Papusus were found in regions where collembolans were

prevalent, but mites were infrequently encountered. Myrmecophily in the Scydmaenidae. Although numerous accounts have been

published about occurrence of scydmaenids in ant nests, there is very little or no information about location of scydmaenids in the colony or about their behavior.

Donisthorpe (1927b:40) published the only observations of scydmaenid behavior when they were associated with ants. He mentioned finding specimens of Stenichnus godarti (Latreille) ". . . in the very centre of the nests (of Formica rufa Linnaeus), running about, evidently quite at home in the midst of the ants, and moving the antennae rapidly in a true myrmecophilous manner". He also collected Eutheia pli cata Gyllenhal in association with F. rufa and observed "the beetle was quite at

home in their (F. rufa) midst; it "twiddled" the antennae very rapidly, vibrating its whole body, and was not attacked when placed in a tube with some of the ants." There are too few published observations of scydmaenid behavior to know whether the actions observed by Donisthorpe are typical of scydmaenid behavior or are found

only among myrmecophilous species. Waller and Moser (1990) infrequently collected a species of Scydmaenus from

nests of Atta texana (Buckley). They found the beetles in detritus cavities (2.5-2.75 m depth), a dormitory cavity (3 m), and a fungus garden (4 m). They did not describe the beetles' behavior or mention how the ants and beetles interacted. Because scyd

maenids are predatory and were found in non-brood chambers, they are probably feeding on other nest associates rather than on ant larvae or eggs. Waller and Moser

(1990) list numerous other myrmecophiles of A. texana, including numerous species of mites, collembolans, thysanurans, roaches, etc.

Franc (1992) and Hlavack and Lackner (1998) listed the known scydmaenid myr mecophiles for Slovakia. They listed Euconnus (Napochus) chrysocomus Saulcy as

a symphillic myrmecophile with Tetramorium caespitum (Linnaeus), E. (N.) claviger (Muller and Kunze), E. (N.) pragensis Machulka, and E. (Neonapochus) maeklini

(Mannerheim) as synoeciosymphillic myrmecophiles, and Euthiconus conicicollis

(Fairmaire and Laboullbene), Neuraphes carinatus (Mulsant), N. imitator Blattny, Scydmoraphes minutus (Chaudoir), Stenichnus foveola Rey, Scydmaenus perrisi Reitter, and S. hellwigi Herbst as synoecious myrmecophiles. Franc (1992) stated



Table 1. Scydmaenid: Ant associations.

Scydmaenidae Formicidae Reference

CLIDICINI O Clidicus formicarius Pascoe Leptogenys diminuta (Smith) Wasmann, 1896:16 C

Clidicus grandis Laporte Leptogenys diminuta (Smith) Wasmann, 1896:16 Z Papusus macer Casey Pogonomyrmex californicus (Buckley) new record >

Pogonomyrmex rugosus Emery new record O CHEVROLATIINI Chevrolatia insignis Jacquelin du Val ants Wasmann, 1894:122 CEPHENNIINI | > Cephennium carpathicum Saulcy Formica rufa Linnaeus Lazorko, 1962:302 t

Lasius brunneus (Latreille) Lazorko, 1962:302 Lasius fuliginosus (Latreille) Lazorko, 1962:302 0

Cephennium majus Reitter Formica rufa Linnaeus Lazorko, 1962:283 0

Formica sp. Koch, 1989:192 Lasius brunneus (Latreille) Lazorko, 1962:283 z Lasius fuliginosus (Latreille) Lazorko, 1962:302 H

Cephennium strudii Besuchet Camponotus ligniperdus (Latreille) Peez, 1957:31 0

EUTHEIINI 0 Eutheia cylindricollis Roubal Fornica rufa Linnaeus Roubal, 1931:68 0 Eutheia formicetorum Reitter Lasius brunneus (Latreille) Donisthorpe, 1927b:40

Myrmica sp. Donisthorpe, 1927b:40 Formica rufa Linnaeus Koch, 1989:191 ants Wasmann, 1894:123; Bernard, 1968:382 0

Eutheia laetificans Roubal Lasius brunneus (Latreille) Roubal, 1931:67 Eutheia linearis Mulsant Formica rufa Linnaeus Koch, 1989:192 H

Lasius brunneus (Latreille) Scriba, 1911:318 Lasius sp. Koch, 1898:192 Myrnica sp. Scriba, 1911:318 o

Eutheia plicata Gyllenhal Formica exsecta Nylander Wasmann, 1894:123; Reitter, 1909:222; Hansen, o 1911:196; Donisthorpe, 1927b:40; Bernard, oo 1968:382; Franz, 1971b:60; Koch, 1989:191



Table 1. Continued.


Formica rufa Linnaeus Wasmann, 1894:123; Everts, 1903:390; Reitter, > 1909:222; Hansen, 1911:196; Donisthorpe, (

1927b:40; Bernard, 1968:382; Franz, 1971b: 0

60

Eutheia scydmaenoides Stephens Formica rufa Linnaeus Wasmann, 1894:123; Everts, 1903:391; Hansen, I3

1911:198 0 z

Lasius fuliginosus (Latreille) Hansen, 1911:198; Roubal, 1931:67 cn

Lasius sp. Koch, 1989:191 O

Euthiconus conicicollis Camponotus vagus (Scopoli) Peez, 1957:31 > (Fairmaire and Laboullbene) Formica rufa Linnaeus Koch, 1989:191, Franc, 1992:305; Hlavac and Z

Lackner, 1998:7

Lasius brunneus (Latreille) Wasmann, 1894:123; Franc, 1992:305; Hlavac

and Lackner, 1998:7

Lasius fuliginosus (Latreille) Hansen, 1911:199; Franc, 1992:305; Hlavac and O

Lackner, 1998:7 Z Lasius sp. Koch, 1989:191

Leptothorax sp. Peez, 1957:31

ants Reitter, 1909:222 H LEPTOMASTACINI i Leptomastax delarouzei Brisout Pheidole sp. Wasmann, 1894:125; Bernard, 1968:383

Leptomastax hypogaeus Pirazzoli Pheidole pallidula (Nylander) Wasmann, 1894:125; Bernard, 1968:383 |

PLAUMANNIOLINI rz Plaumanniola sanctaecatharinae Costa Lima Octostruma stenognatha Brown and Kempf Lenko 1972:327; Franz, 1990:32 0

SCYDMAENINI Ceramphis deformata (Horn) Solenopsis aurea Wheeler O'Keefe, 1997b:342

Palaeoscydmaenus australis Franz Pachycondyla lutea (Mayr) Franz, 1975b:273

Scydmaenus abbreviatellus Erichson Formica rufa Linnaeus Markel, 1845:242

Scydmaenus angulatus Muller and Kunze Formica conicularia Latreille Markel, 1845:242

Scydmaenus bifasciculatus Lea Rhytidoponera metallica (Smith) Lea, 1912:57



Table 1. Continued.


Scydmaenus carinatus (King) Amblyopone australis Erichson Lea, 1912:60 Papyrius nitidus (Mayr) Lea, 1910:187 ants Wasmann, 1894:125: Lea, 1905:377 Z

Scydmaenus castaneoglaber Lea Camponotus gasseri (Forel) Lea, 1910:185 >

Rhytidoponera metallica (Smith) Lea, 1912:60 O Scydmaenus cerastes Baudi Myrmica rubra (Linnaeus) Marsuel, 1872:41 T

Scydmaenus clarkianus Franz Pachycondyla lutea (Mayr) Franz, 1975b:279

Scydmaenus clavatus LeConte Crematogaster lineolata (Say) Wasmann, 1894:124 z Scydmaenus colobopsis Lea Amblyopone australis Erichson Lea, 1912:59

Camponotus gasseri (Forel) Lea, 1910:182 Scydmaenus cornutus Motsch. Messor sp. Wasmann, 1894:125 |

Myrmica cephalotes Motschulsky, 1851:503 Scydmaenus daveyi Lea Camponotus sp. Lea, 1910:182 Scydmaenus duplicates Lea ants Lea, 1910:184 Z Scydmaenus ectatommae Lea Rhytidoponera metallica (Smith) Lea, 1910:183 0O

Scydmaenus formnicetorum Franz Camponotus rufipes (Fabricius) Franz, 1980b:54 0

Scydmaenus formicarum Franz Lasius sp. Franz, 1975b:277

Scydmaenus hellwigi Herbst Formnica rufa Linnaeus Motschulsky, 1851:503; Reitter, 1882:198; Was- 0

mann, 1894:124; Everts, 1903:395; Reitter,

1909:228; Hansen, 1911:220; Bernard, 1968: 2

382; Koch, 1989:200; Franc, 1992:307;

Drozd, 1995:386; Hlavac and Lackner, 1998:7 0

Lasius brunneus (Latreille) Wasmann, 1894:124; Everts, 1903:395; Peez, 1957:31; Bernard, 1968:382; Franc, 1992:307; H

Drozd, 1995:386; Hlavac and Lackner, 1998: <

3, 7

Lasius fuliginosus (Latreille) Wasmann, 1894:124; Everts, 1903:395; Bernard, o 1968:382; Franc, 1992:307; Drozd, 1995:386; o

Hlavac and Lackner, 1998:7 oo

Lasius sp. Reitter, 1909:228; Koch, 1989:200



Table 1. Continued.


Scydmaenus hirticollis Gyllenhal Formica conicularia Latreille Markel, 1845:242 Scydmaenus impavidus Lea ants Lea, 1912:56 (A Scydmaenus incerticornis Lea Aphaenogaster longiceps (Smith) Lea, 1912:58 O

Pachycondyla lutea (Mayr) Lea, 1912:58 Scydmaenus incola Franz Camponotus rufipes (Fabricius) Franz, 1980b:43 H Scydmaenus incolaeformis Franz Odontomachus affinis Guerin-Meneville Franz, 1980b:43 O Scydmaenus lenkoanus Franz Camponotus rufipes (Fabricius) Franz, 1980b:56 z Scydmaenus longicollis Machulka Fonnica rufa Linnaeus Machulka, 1931:85 O Scydmaenus microps Lea Pachycondyla lutea (Mayr) Lea, 1910:185 T1 Scydmaenus myrmecophilus Menozzi Messor structor (Latreille) Menozzi, 1941:6 > Scydmaenus optatus Sharp Camponotus dorycus (Smith) Lea, 1912:186 H

Ectatomma melaleuca Franz, 1975b:281 Pachycondyla lutea (Mayr) Lea, 1910:186 Iridomyrmex conifera (Forel) Clark, 1920:102

Scydmaenus perrisi Reitter Lasius brunneus (Latreille) Reitter, 1909:228; Meybohm, 1971:102; Franc, 0 1992:306; Hlavac and Lackner, 1998:7 z

Lasius sp. Koch, 1989:200 m

ants Wasmann, 1894:125 t'1 Scydmaenus pinguiculus Gerstaeker Messor sp. Gerstaecker, 1873:84 H Scydmaenus rufus Muller and Kunze Formica rufa Linnaeus Wasmann, 1894:124: Everts, 1903:395; Reitter, t

1909:228; Bernard, 1968:382

Lasius brunneus (Latreille) Reitter, 1909:228 O Lasiusfuliginosus (Latreille) Everts, 1903:395 r Myrmica sp. Peez, 1957:31 1

Scydmaenus simplicicornis Lea Papyrius nitidus (Mayr) Lea, 1910:186 Scydmaenus swanensis Franz Pachycondyla lutea (Mayr) Franz, 1975b:281 Scydmaenus vanboveni Franz Paratrechina sp. Franz, 1976:971 > Scydmaenus sp. Atta texana (Buckley) Waller and Moser, 1990:266

CYRTOSCYDMAENINI Alloraphes lenkoi Franz Camponotus rufipes (Fabricius) Franz, 1980b:215 Alloraphes myrmecophilus Franz Camponotus rufipes (Fabricius) Franz, 1980b:214



Table 1. Continued.


Euconnus O (Dimorphoconnus) insigniventris (Lea) Rhytidoponera metallica (Smith) Lea, 1912:59; Franz, 1975b:178

(Euconophron) clientulus (Lea) Rhytidoponera metallica (Smith) Lea, 1910:181; Franz, 1975b:271 Z

(Euconophron) glabripennis (Lea) Polyrhachis hexancantha (Erichson) Lea, 1910:183; Franz, 1975b:223

(Euconophron) lucindalensis Franz Rhytidoponera metallica (Smith) Franz, 1975b:215 0

(Napochus) balaena Saulcy Tetramorium caespitum (Linnaeus) Escherich, 1897:233 H

(Napochus) camponotorum Franz Camponotus rufipes (Fabricius) Franz, 1980b: 102

(Napochus) chrysocomus Saulcy Messor barbarus (Linnaeus) Reitter, 1909:226; Binaghi, 1956:73; Porta, z

1959:148; Koch, 1989:198

Messor structor (Latreille) Koch, 1989:198

Tetramorium caespitum (Linnaeus) Wasmann, 1894:124; Escherich, 1897:233; Reit- O

ter, 1909:226; Binaghi, 1956:73; Peez, 1957: t

31; Porta, 1959:148; Bernard, 1968:382;

Koch, 1989:198; Franc, 1992:306; Hlavac and z

Lackner, 1998:3, 7 0

(Napochus) clavicorne (Casey) ants Blatchley, 1910:299 0

(Napochus) claviger (Muller and Kunze) Camponotus ligniperdus (Latreille) Peez, 1957:31

Formica fusca Linnaeus Machulka, 1928:70; Binaghi, 1956:74; Peez, 0

1957:31 Formica rufa Linnaeus Reitter, 1882:181; Wasmann, 1894:124; Everts, :

1903:394; Reitter, 1909:226; Hansen, 1911: 0

214; Donisthorpe, 1926:150; Machulka, 1923: n

70; Donisthorpe, 1927b:40; Binaghi, 1956:74; tt

Peez, 1957:31; Porta, 1959:148; Bernard, H

1968:382; Meybohm, 1971:101; Koch, 1989:

198; Franc, 1992:306; Hlavac and Lackner,

1998:7 Formica sanguinea (Latreille) Machulka, 1923:70; Binaghi, 1956:74; Porta,

1959:148



Table 1. Continued.


Lasius brunneus (Latreille) Donisthorpe 1926:150; Donisthorpe 1927b:40; > Nohel 1970:136; Hlavac and Lackner 1998:7 m

Lasius fuliginosus (Latreille) Wasmann, 1894:124; Everts, 1903:394; Reitter, 0

1909:226; Hansen, 1911:214; Donisthorpe,

1926:150; Machulka, 1923:70; Donisthorpe, 3 1927b:40; Binaghi, 1956:74; Peez, 1957:31; 0

z Porta, 1959:148; Bernard, 1968:382; Hlavac CA

and Lackner, 1998:7 O Lasius niger (Linnaeus) Wasmann, 1894:124; Everts, 1903:394; Reitter, >

1909:226; Hansen, 1911:214; Donisthorpe, z 1926:150; Machulka, 1923:70; Binaghi, 1956: -

74; Porta, 1959:148; Bernard, 1968:382; Mey

bohm, 1971: 101

Lasius sp. Koch, 1989:198 0 ants Miirkel, 1845:242 z

(Napochus) cornutus (Saulcy) Messor capitatus (Latreille) Saulcy, 1863:655 t

Messor barbarus (Linnaeus) Saulcy, 1864:257; Wasmann, 1894:124; Bernard, t

1968:382 H Messor sp. Binaghi, 1956:76; Porta, 1959:148

(Napochus) fulvum (LeConte) ants Blatchley, 1910:300 (Napochus) globiceps Franz Camponotus sp. Franz, 1980b: 117 0 (Napochus) lenkoi Franz Camponotus rufipes (Fabricius) Franz, 1880b:100

Odontomachus affinis Guerin-Meneville Franz, 1980b: 100 O (Napochus) odontomachi Franz Odontomachus affinis Guerin-Meneville Franz, 1980b:82 (Napochus) paracamponotorum Franz Camponotus rufipes (Fabricius) Franz, 1980b: 104 (Napochus) paulinus Franz Camponotus rufipes (Fabricius) Franz, 1980b:94 (Napochus) pragensis Machulka Formica rufa Linnaeus Peez, 1957:31; Franc, 1992:306

Lasius brunneus (Latreille) Machulka, 1923:70; Binaghi, 1956:76; Nohel,

1970:136; Franc, 1992:306 |



Table 1. Continued. |


Lasius fuliginosus (Latreille) Hlavac and Lackner, 1998:7

Lasius sp. Koch, 1989:199 Myrmica rubra (Fabricius) Peez, 1957:31 z

(Napochus) rufobrunneus Franz Camponotus rufipes (Fabricius) Franz, 1980b:87 >

(Napochus) saopauli Franz Camponotus rufipes (Fabricius) Franz, 1980b:81 0

(Neonapochus) maeklini (Mannerheim) Formica rufa Linnaeus Mannerheim, 1844:193; Wasmann, 1894:124; Everts, 1903:394; Reitter, 1909:226; Binaghi, Z 1956:78; Koch, 1989:199; Franc, 1992:306; m

Drozd, 1995:386; Hlavac and Lackner, 1998:7 t

Lasius brunneus (Latreille) Meybohm, 1971:101; Franc, 1992:306; Drozd, X 1995:386; Hlavac and Lackner, 1998:7 0

Lasius fuliginosus (Latreille) Franc, 1992:306; Drozd, 1995:386; Hlavac and O

Lackner, 1998:7

Lasius sp. Koch, 1989:199 z

(Pycnophus) nidicola Franz Camponotus rufipes (Fabricius) Franz, 1980b: 118

(Pycnophus) paranidicola Franz Camponotus rufipes (Fabricius) Franz, 1993:48 g

(Pycnophus) verticicoais Franz Camponotus rufipes (Fabricius) Franz, 1980b:4122

Euconnus baruerii Franz Camponotus rufipes (Fabricius) Franz, 1980b: 172 0

Euconnus denticornis (Muller) Formica rufa Linnaeus Hansen, 1911:215

Euconnus lenkoanus Franz Camponotus rufipes (Fabricius) Franz, 1980b: 161

Euconnus semiruber Casey ants Blatchely, 1910:294 Microscydmus myrmecophilus Franz Camponotus rufipes (Fabricius) Franz, 1980b:203 0

Microscydmus nanus (Schaum) Camponotus ligniperdus (Latreille) Peez, 1957:31

Camponotus vagus (Scopoli) Peez, 1957:31 H Formica rufa Linnaeus Hansen, 1911:217

Formica sp. Koch, 1989:198 Leptothorax sp. Peez, 1957:31 Lasius sp. Koch, 1989:198

00 Mimoscydmus baruerri Franz Camponotus rufipes (Fabricius) Franz, 1980b:220

Napochomorphus extraneus Franz Camponotus rufipes (Fabricius) Franz, 1980b:206



Table 1. Continued.


Neuraphes balaena Saulcy Tetramorium caespitum (Linnaeus) Franz, 1975:16 > Neuraphes besucheti Franz Formica rufa Linnaeus Franz, 1966a:233 W

Neuraphes carinatus (Mulsant) Formica rufa Linnaeus Meybohm, 1971:100; Koch, 1989:194; Hlavac O

and Lackner, 1998:7

Lasius brunneus (Latreille) Machulka, 1931:81; Koch, 1989:194; Hlavac >

and Lackner, 1998:7 0 Myrmica sp. Franc, 1992:305 Tetramorium sp. Koch, 1989:194 0

Neuraphes coecus Reitter Lasius brunneus (Latreille) Koch, 1989:196

Neuraphes elongatus (Muller and Kunze) Formica conicularia Latreille Miirkel, 1845:242 z

Formica rufa Linnaeus Hansen, 1911:205 H Neuraphes imitator Blattny Formica sp. Koch, 1989:194

Lasius brunneus (Latreille) Machulka, 1931:81 Lasius sp. Koch, 1989:194 H Ponera coarctata (Latreille) Hlavac and Lackner, 1998:7 0

Ponera sp. Koch, 1989:194 Neuraphes myrmecophilus Aube ants Wasmann, 1894:123

Neuraphes parvulus Rambousek Lasius fuliginosus (Latreille) Rambousek, 1909:17

Neuraphes plicicollis Reitter Formica polyctena Foerster Vorst, 1988:162 H

Neuraphes rubicundus Schaum Formica conicularia Latreille Markel, 1845:242 z

Formica rufa Linnaeus Hansen, 1911:203; Machulka, 1931:79

Lasius fuliginosus (Latreille) Hansen, 1911:203; Machulka, 1931:80

Neuraphes slovenicus Machulka Camponotus aethiops (Latreille) Machulka, 1931:83 0

Neuraphes sparshalli (Denny) Lasius fuliginosus (Latreille) Wasmann, 1894:123; Everts, 1903:392; Hansen, ~e

1911:207 tHi Neuraphes talparum Lokay ants Machulka, 1931:80

Scydmoraphes geticus (Saulcy) Formica rufa Linnaeus Machulka, 1931:89; Seriani, 1987:248

Formica sp. Koch, 1989:196 Lasius brunneus (Latreille) Machulka, 1931:89; Peez, 1957:31; Seriani,

1987:248 0 ., _ s~~~~~~



Table 1. Continued. |0


Lasius sp. Koch, 1989:196 Lasius fuliginosus (Latreille) Machulka, 1931:89; Seriani, 1987:248

Scydmoraphes menozzii Franz Messor structor (Latreille) Franz, 1966b:570 z Scydmoraphes minutus (Chaudoir) Camponotus vagus (Scopoli) Peez, 1957:31 >

Formica rufa Linnaeus Reitter, 1882:172; Wasmann, 1894:123; Everts, O 1903:393; Reitter, 1909:225; Hansen, 1911:

219; Machulka, 1931:89; Bernard, 1968:382; =

Koch, 1989:196 z Lasius brunneus (Latreille) Machulka, 1931:89

Lasius fuliginosus (Latreille) Reitter, 1909:225; Machulka, 1931:89 Lasius sp. Koch, 1989:196

0 Leptothorax sp. Peez, 1957:31 x Stenichnus bicolor (Denny) Formica rufa Linnaeus Koch, 1989:197 Stenichnus collaris (Muller and Kunze) Formica conicularia Latreille Markel, 1845:242 z

Formica rufa Linnaeus Markel, 1945:242; Hansen, 1911:212 H Lasius fuliginosus (Latreille) Markel, 1845:242; Hansen, 1911:212

Stenichnus compendiensis Mequignon Lasius brunneus (Latreille) Mequignon, 1907:218 r

Stenichnus constricta (Lea) Amblyopone australis Erichson Lea, 1910:187 0 Stenichnus exilis (Erichson) Formica exsecta Nylander Wasmann, 1894:123; Everts, 1903:393; Donis- C!

thorpe, 1927b:40 Formica rufa Linnaeus Wasmann, 1894:123; Everts, 1903:393; Donis- r

thorpe, 1927b:40 0 Lasius brunneus (Latreille) Donisthorpe, 1927b:40; Nohel, 1970:136

Stenichnus foveola Rey Formica sp. Koch, 1989:197 H Lasius sp. Koch, 1989:197 ants Franc, 1992:306

Stenichnus godarti (Latreille) Camponotus herculeanus (Linnaeus) Nohel, 1970:136 Formica rufa Linnaeus Markel, 1845:242; Kraatz, 1865:111; Mequeg- o

non, 1907:218; Hansen, 1911:210; Donis

thorpe, 1927b:40; Koch, 1989:197



O

Table 1. Continued. c 0


Lasius brunneus (Latreille) Donisthorpe, 1927b:40; Nohel, 1970:136 H 0

Lasius fuliginosus (Latreille) Markel, 1845:242; Everts, 1903:393; Hansen, z

1911:210; Franz, 1971a:71

Lasius sp. Koch, 1989:197 Stenichnus peezi Franz Lasius brunneus (Latreille) Koch, 1989:196 Stenichnus pelliceus Holdhaus Tetramorium caespitum (Linnaeus) Koch, 1989:197 H Stenichnus pusilla (King) Aphaenogaster longiceps (Smith) Lea, 1912:60

Monomorium sp. Lea, 1912:60 Pachycondyla lutea (Mayr) Lea, 1910:187; Lea 1912:60

Rhytidoponera metallica (Smith) Lea, 1912:60

Stigmacros froggatti (Forel) Lea, 1912:60 Z Stenichnus pusillum Muller and Kunze Formica rufa Linnaeus Koch, 1989:197

Stenichnus scutellaris (Muller and Kunze) Formica conucularia Latreille Markel, 1845:242

Formica rufa Linnaeus Hansen, 1911:211; Peez, 31 H Lasius fuliginosus (Latreille) Everts, 1903:393; Hansen, 1911:211 cr1

Stenichnus stotti Donisthorpe Lasius fuliginosus (Latreille) Donisthorpe, 1932:268

Myrmecina graminicola (Latreille) Donisthorpe, 1932:267 0

Stenichnus syriacus Franz Lasius fuliginosus (Latreille) Koch, 1989:197

Syndicus kingi King Iridomyrmex rufoniger (Lowne) Lea, 1910:188 O

Syndicus latipennis Lea ants Lea, 1910:188 H Syndicus macrosticta Lea ants (Iridomyrmex?) Lea, 1910:189

00



Table 2. Ant: Scydmaenid associations. |0

Formicidae Scydmaenidae Reference

PONERINAE O Amblyopone australis Erichson Heterognathous myrmecophilus Lea Lea, 1912:60

Stenichnus constricta (Lea) Lea, 1910:187 z Scydmaenus colobopsis Lea Lea, 1912:59 >

Ectatomma melaleuca Scydmaenus optatus Sharp Franz, 1975b:281 0 Leptogenys diminuta (Smith) Clidicus formicarius Pascoe Wasmann, 1896:16

Clidicus grandis Laporte Wasmann, 1896:16 Odontomachus affinis Guerin-Meneville Euconnus (Napochus) lenkoi Franz Franz, 1980b: 100

Euconnus (Napochus) odontomachi Franz Franz, 1980b:82 z

Scydmaenus incolaeformis Franz Franz, 1980b:43 Pachycondyla lutea (Mayr) Palaeoscydmaenus australis Franz Franz, 1975b:273

Stenichnus pusilla (King) Lea, 1910:187; Lea, 1912:60 P Scydmaenus incerticornis Lea Lea, 1912:60 Scydmaenus microps Lea Lea, 1910:185 z Scydmaenus clarkianus Franz Franz, 1975b:279 H Scydmaenus optatus Sharp Lea, 1910:186

0

Scydmaenus swanensis Franz Franz, 1975b:281 0 Ponera coarctata (Latreille) Neuraphes imitator Blattny Hlavack and Lackner, 1998:7 0 Ponera sp. Neuraphes imitator Blattny Koch, 1989:194 Q

Rhytidoponera metallica (Smith) Euconnus (Dimorphoconnus) insigniventris (Lea) Lea, 1912:59; Franz, 1975b:178 Euconnus (Euconophron) clientulus (Lea) Lea, 1910:181; Franz, 1975b:271 Euconnus (Euconophron) lucindalensis Franz Franz, 1975b:215 0 Stenichnus pusilla (King) Lea, 1912:60

Scydmaenus bifasciculatus Lea Lea, 1912:57 H Scydmaenus castaneoglaber Lea Lea, 1912:60

Scydmaenus ectatommae Lea Lea, 1910:183

MYRMICINAE Aphaenogaster longiceps (Smith) Stenichnus pusilla (King) Lea, 1912:60

Scydmaenus incerticornis Lea Lea, 1912:58 Atta texana (Buckley) Scydmaenus sp. Waller and Moser, 1990:266



Table 2. Continued. 0


Crematogaster lineolata (Say) Scydmaenus clavatus LeConte Wasmann, 1894:281 > Leptothorax sp. Microscydmus nanus (Schaum) Peez, 1957:31 Cn

Euthiconus conicicollis Peez, 1957:31 O (Fairmaire and Laboullbene)

Scydmoraphes minutus (Chaudoir) Peez, 1957:31 >

Messor barbarus (Linnaeus) Euconnus (Napochus) chrysocomus Saulcy Reitter, 1909:226; Binaghi, 1956:73; Porta, O z 1959:148; Koch, 1989:198 z

Euconnus (Napochus) cornutus Saulcy Wasmann, 1894:124; Bernard, 1968:382 0 Messor capitatus (Latreille) Euconnus (Napochus) cornutus Saulcy Saulcy, 1863:655

Messor structor (Latreille) Euconnus (Napochus) chrysocomus Saulcy Hlavack and Lackner, 1998:7 z

Scydmaenus myrmecophilus Menozzi Menozzi, 1941:6 H Scydmoraphes menozzii Franz Franz, 1966b:570

Messor spp. Euconnus (Napochus) cornutus Saulcy Binaghi, 1956:76; Porta, 1959:148

Scydmaenus pinguiculus Gerstaecker Gerstaecker, 1873:84 H Monomorium sp. Stenichnus pusilla (King) Lea, 1912:60 O

Myrmecina graminicola (Latreille) Stenichnus stotti Donisthorpe Donisthorpe, 1932:267 tIl

Octostruma stenognatha Brown and Kempf Plaumanniola sanctaecatharinae Costa Lima Lenko, 1972:327; Franz, 1990:32

Pheidole pallidula (Nylander) Leptomastax hypoganaeus Pirazzoli Wasmann, 1894:125; Bernard, 1968:383 t

Pheidole sp. Leptomastax delarouzei Brisout Wasmann, 1894:125; Bernard, 1968:383 H

Pogonomyrmex californicus (Buckley) Papusus macer Casey new record v Pogonomyrmex rugosus Emery Papusus macer Casey new record

Solenopsis aurea Wheeler Ceramphis deformata (Horn) O'Keefe, 1997b:342

Tetramorium caespitum (Linnaeus) Euconnus (Napochus) chrysocomus Saulcy Wasmann, 1894:124; Escherich, 1897:233; O 0

Reitter, 1909:226; Binaghi, 1956:73; Peez, 1957:31; Porta, 1959:148; Bernard, H

1968:382; Koch, 1989:198; Franc, 1992:

306; Hlavac and Lackner, 1998:3, 7

Neuraphes balaena (Saulcy) Escherich, 1897:233; Franz, 1975:16

Stenichnus pelliceus Holdhaus Koch, 1989:197 Tetramorium sp. Neuraphes carinatum Mulsant Koch, 1989:194 0

* _ .. . . ..~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~



w Table 2. Continued. 00

oo


DOLICHODERINAE O

Iridomyrmex conifera (Forel) Scydmaenus opatus Sharp Clark, 1920:102

Iridomyrmex rufoniger (Lowne) Syndicus kingi King Lea, 1910:188

Iridomyrmex? Syndicus macrostricta (Lea) Lea, 1910:189 Papyrius nitidus (Mayr) Heterognathus carinatus King Lea, 1910:187 0

Scydmaenus simplicicornis Lea Lea, 1910:186 H FORMICINAE Camponotus aethiops (Latreille) Neuraphes slovenicus Machulka Machulka, 1931:83 z

Camponotus dorycus (Smith) Scydmaenus optatus Sharp Lea, 1910:186

Camponotus gasseri (Forel) Scydmaenus castaneoglaber Lea Lea, 1910:185

Scydmaenus colobopsis Lea Lea, 1910:182 O Camponotus herculeanus (Linnaeus) Stenichnus godarti (Latreille) Nohel, 1970:136

Camponotus ligniperdus (Latreille) Cephennium strudii Besuchet Peez, 1957:31 tr Euconnus (Napochus) claviger (Muller) Peez, 1957:31 z

Microscydmus nanus (Schaum) Peez, 1957:31 0

Camponotus rufipes (Fabricius) Alloraphes lenkio Franz Franz, 1980b:215 o

Alloraphes myrmecophilus Franz Franz, 1980b:214 O Euconnus baruerii Franz Franz, 1980b: 172 Euconnus lenkoanus Franz Franz, 1980b: 161 ) Euconnus (Napochus) camponotorum Franz Franz, 1980b: 102

Euconnus (Napochus) lenkoi Franz Franz, 1980b: 100 0 Euconnus (Napochus) paracamponotorum Franz Franz, 1980b: 104 Q Euconnus (Napochus) paulinus Franz Franz, 1980b:94 H Euconnus (Napochus) ruffobrunneus Franz Franz, 1980b:87 Euconnus (Napochus) saopauli Franz Franz, 1980b:81

Euconnus (Pycnophus) paranidicola Franz Franz, 1993:48 o

Euconnus (Pycnophus) nidicola Franz Franz, 1980b: 118

Euconnus (Pycnophus) verticicornis Franz Franz, 1980b: 112 oo Microscydmus myrmecophilus Franz Franz, 1980b:203 -IL



Table 2. Continued.


Mimoscydmus baruerii Franz Franz, 1980b:220 > Napochomorphus extraneus Franz Franz, 1980b:206 (A

Scydmaenus formicitorum Franz Franz, 1980b:54 0 Scydmaenus incola Franz Franz, 1980b:43 Scydmaenus lenkoanus Franz Franz, 1980b:56

Camponotus vagus (Scopoli) Euthiconus conicicollis Peez, 1957:31 O

(Fairmaire and Laboullbene) (A

Microscydmus nanus (Schaum) Peez, 1957:31 O

Scydmoraphes minutus (Chaudoir) Peez, 1957:31 > Camponotus spp. Euconnus (Napochus) globiceps Franz Franz, 1980:117 Z

Scydmaenus daveyi Lea Lea, 1910:182 Formica cunicularia Latreille Neuraphes elongatus (Muller and Kunze) Markel, 1845:242

Neuraphes rubicundus Schaum Markel, 1845:242

Scydmaenus angulatus Muller and Kunze Miirkel, 1845:242 0 Scydmaenus hirticollis Gyllenhal Miirkel, 1845:242 Z Stenichnus collaris (Muller and Kunze) Markel, 1845:242 t

Stenichnus scutellaris Muller and Kunze Mairkel, 1845:242 Formica exsecta Nylander Eutheia plicata Gyllenhal Wasmann, 1894:123; Reitter, 1909:222; r

Hansen, 1911:196; Donisthorpe, 1927b: tr

40; Bernard, 1968:382; Franz, 1971b:

60 O Stenichnus exilis (Erichson) Wasmann, 1894:123; Everts, 1903:393;

Donisthorpe, 1927b:40 |

Formica fusca Linnaeus Euconnus (Napochus) claviger (Muller) Machulka, 1923:70; Binaghi, 1956:74; Por- H ta, 1959:148

Formica polyctena Foerster Neuraphes plicicollis Reitter Vorst, 1988:162 Formica rufa Linnaeus Cephennium carpathicum Saulcy Lazorko, 1962:302

Cephennium majus Reitter Lazorko, 1962:283 Euconnus denticornis (Muller) Hansen, 1911:215 s



Table 2. Continued.


Euconnus (Napochus) claviger (Muller) Reitter, 1882:181; Wasmann, 1894:124;

Everts, 1903:394; Reitter, 1909:226; Han- C

sen, 1911:214; Donisthorpe, 1926:150; z

Machulka, 1923:70; Donisthorpe, 1927b: >

40; Binaghi, 1956:74; Peez, 1957:31; O

Porta, 1959:148; Bernard, 1968:382;

Meybohm, 1971:101; Koch, 1989:198;

Franc, 1992:306; Hlavack and Lackner,

1998:7 Euconnus (Napochus) pragensis Machulka Peez, 1957:31; Franc, 1992:306

Euconnus (Neonapochus) maeklini (Mannerheim) Mannerheim, 1844:193; Wasmann, 1894: 0

124; Everts, 1903:394; Reitter, 1909:226; P

Binaghi, 1956:78; Bernard, 1968:382;

Koch, 1989:199; Franc, 1992:306; Drozd, z

1995:386; Hlavack and Lackner, 1998:7 O

Eutheia cylindricollis Roubal Roubal, 1931:68 0

Eutheia formicetorum Reitter Koch, 1989:191 r

Eutheia linearis Mulsant Koch, 1989:192 0 Eutheia plicata Gyllenhal Wasmann, 1894:123; Everts, 1903:390;

Reitter, 1909:222; Hansen, 1911:196;

Donisthorpe, 1927b:40; Bernard, 1968: r

382; Franz, 1971b:60; Koch, 1989:191 0

Eutheia scydmaenoides Stephens Wasmann, 1894:123; Everts, 1903:391;

Hansen, 1911:198 H Euthiconus conicicollis Franc, 1992:305; Koch, 1989:191; Hlavack

(Fairmaire and Laboullbene) and Lackner, 1998:7

Microscydmus nanus (Schaum) Hansen, 1911:217

Neuraphes besucheti Franz Franz, 1966a:233 o

Neuraphes carinatus Mulsant Meybohm, 1971:100; Koch, 1989:194; Hla- o

vack and Lackner, 1998:7



Table 2. Continued. 8


Neuraphes elongatus (Muller and Kunze) Hansen, 1911:205 > Neuraphes rubicundus Schaum Hansen, 1911:203; Machulka, 1931:79 CA

Scydmaenus abbreviatellus Erichson Markel, 1845:242 0 Scydmaenus longicollis Motschulsky Motschulsky, 1845:49; Machulka, 1931:85 n

Scydmaenus rufus Muller and Kunze Wasmann, 1894:124; Everts, 1903:395; H

Reitter, 1909:228; Bernard, 1968:382 O

Scydmaenus hellwigi Herbst Motschulsky, 1851:503; Reitter, 1882:198; Z

Wasmann, 1894:124; Everts, 1903:395; 0

Reitter, 1909:228; Hansen, 1911:220; Bernard, 1968:382; Koch, 1989:200; z Franc, 1992:307; Drozd, 1995:386; Hla


Scydmoraphes geticus Saulcy Machulka, 1931:89; Seriani, 1987:248

Scydmoraphes minutus (Chaudoir) Reitter, 1882:172; Wasmann, 1894:123; <

Everts, 1903:393; Reitter, 1909:225; Han- 0 z

sen, 1911:219; Machulka, 1931:89; Ber- tx

nard, 1968:382; Koch, 1989:196; Hla

vack and Lackner, 1998:7 H Stenichnus bicolor (Denny) Koch, 1989:197

Stenichnus collaris (Muller and Kunze) Markel, 1845:242; Hansen, 1911:212 m

Stenichnus exilis (Erichson) Wasmann, 1894:123; Everts, 1903:393;

Donisthorpe, 1927b:40 r

Stenichnus.foveola Rey Koch, 1989:197 0 Stenichnus godarti (Latreille) Markel, 1845:242; Kraatz, 1865:111; Me- ~ e

quegnon, 1907:218; Hansen, 1911:210; tT

Donisthorpe, 1927b:40; Koch, 1989:197 > Stenichnus pusillus Muller and Kunze Koch, 1989:197

Stenichnus scutellaris (Muller and Kunze) Hansen, 1911:211; Peez, 1957:31

Formica sanguinea (Latreille) Euconnus (Napochus) claviger (Muller) Machulka, 1923:70; Binaghi, 1956:74; Por

ta, 1959:148



Table 2. Continued.


Lasius alienus (Foerster) Euconnus (Pycnophus) rasus (LeConte) Wasmann, 1894:124 O

Lasius brunneus (Latreille) Cephennium carpathicum Saulcy Lazorko, 1962:302

Cephennium majus Reitter Lazorko, 1962:283 z Euconnus (Napochus) claviger (Muller) Donisthorpe, 1926:150; Donisthorpe, >

1927b:40; Nohel, 1970:136; Hlavack and 0

Lackner, 1998:7

Euconnus (Napochus) pragensis Machulka Machulka, 1923:70; Binaghi, 1956:76; No- =

hel, 1970:136; Franc, 1992:306; Hlavack trr z

and Lackner, 1998:7 tT

Euconnus (Neonapochus) maeklini (Mannerheim) Meybohm, 1971:101; Franc, 1992:306;

Drozd, 1995:386; Hlavack and Lackner, O 0 1998:7 O

Eutheia formicetorum Reitter Donisthorpe, 1927b:40 P Eutheia laetificans Roubal Roubal, 1931:67 z

Eutheia linearis Mulsant Scriba, 1911:318 H

Euthiconus conicicollis Wasmann, 1894:123; Franc, 1992:305; Hla- 0

(Fairmaire and Laboullbene) vack and Lackner, 1998:7 0

Neuraphes carinatus Mulsant Machulka, 1931:81; Koch, 1989:194; Hla- 0


Neuraphes imitator Blattny Machulka, 1931:81

Scydmaenus hellwigi Herbst Wasmann, 1894:124; Everts, 1903:395;

Peez, 1957:31; Bernard, 1968:382; Franc, 0

1992:307; Drozd, 1995:386; Hlavac and Q

Lackner, 1998:3, 7 H

Scydmaenus perrisi Reitter Reitter, 1909:228; Meybohm, 1971:102;

Franc, 1992:306; Hlavack and Lackner, <

1998:7 Scydmaenus rufus Muller and Kunze Reitter, 1909:228

Scydmoraphes geticus (Saulcy) Machulka, 1931:89; Peez, 1957:31; Seriani, oo

1987:248



Table 2. Continued.


Scydmoraphes minutus (Chaudoir) Machulka, 1931:89; Hlavack and Lackner, > Stenichnus compendiansis Mequegnon 1998:7 cn

Stenichnus exilis (Erichson) Mequegnon, 1907:218 O

Stenichnus foveola Rey Donisthorpe, 1927b:40; Nohel, 1970:136 Stenichnus godarti (Latreille) Koch, 1989:197; Hlavack and Lackner,

1998:7 O Donisthorpe, 1927b:40; Nohel, 1970:136 Z

Stenichnus peezi Franz Koch, 1989:196 0 Lasius fuliginosus (Latreille) Cephennium carpathicum Saulcy Lazorko, 1962:302

Cephennium majus Reitter Lazorko, 1962:283 z Euconnus (Napochus) claviger (Muller) Wasmann, 1894:124; Everts, 1903:394;

Reitter, 1909:226; Hansen, 1911:214;

Donisthorpe, 1926:150; Machulka, 1923:

70; Donisthorpe, 1927b:40; Binaghi,

1956:74; Peez, 1957:31; Porta, 1959:148; Z z

Bernard, 1968:382; Hlavack and Lackner, tri

1998:7 t tTl Euconnus (Napochus) pragensis Machulka Hlavack and Lackner, 1998:7

Euconnus (Neonapochus) maeklini (Mannerheim) Franc, 1992:306; Drozd, 1995: 386; Hla

vack and Lackner, 1998:7 tr

Eutheia scydmaenoides Stephens Hansen, 1911:198; Roubal, 1931:67 Euthiconus conicicollis Hansen, 1911:199; Franc, 1992:305; Hla- r

(Fairmaire and Laboullbene) vack and Lackner, 1998:7 0 Neuraphes parvulus Rambousek Rambousek, 1909:17

0

Neuraphes rubicundus Schaum Hansen, 1911:203; Machulka, 1931:80 trr Neuraphes sparshalli (Denny) Wasmann, 1894:123; Everts, 1903:392;

Hansen, 1911:207

Scydmaenus hellwigi Herbst Wasmann, 1894:124; Everts, 1903:395; Ber

nard, 1968:382; Franc, 1992:307; Drozd,

1995:386; Hlavack and Lackner, 1998:7 o



Table 2. Continued. \0


Scvdmaenus rufus Muller and Kunze Everts, 1903:395 O

Scydmoraphes geticus Saulcy Machulka, 1931:89; Seriani, 1987:248

Scydmoraphes minutus (Chaudoir) Reitter, 1909:225; Machulka, 1931:89; Hla- z vack and Lackner, 1998:7 r

Stenichnus collaris (Latreille) Markel, 1845:242; Hansen, 1911:212 O

Stenichnus godarti (Latreille) Miirkel, 1845:242; Everts, 1903:393; Han- 3 sen, 1911:210; Franz, 1971a:71

Stenichnus scutellaris (Muller and Kunze) Everts, 1903:393; Hansen, 1911:211

Stenichnus stotti Donisthorpe Donisthorpe, 1932:268 Stenichnus styriacus Franz Koch, 1989:197

Lasius niger (Linnaeus) Euconnus (Napochus) claviger (Muller) Wasmann, 1894:124; Everts, 1903:394;

Reitter, 1909:226; Hansen, 1911:214;

Donisthorpe, 1926:150; Machulka, 1923: z

70; Binaghi, 1956:74; Porta, 1959:148; z

Bernard, 1968:382; Meybohm, 1971:101; H

Hlavack and Lackner, 1998:7

Lasius sp. Scydmaenus formicarum Franz Franz, 1975:277

Scydmaenus hellwigi Herbst Reitter, 1909:228 0 Paratrechina sp. Scydmaenus vanboveni Franz Franz, 1976:971

Polyrhachis hexacantha (Erichson) Euconnus (Euconnophron) glabripennis (Lea) Lea, 1910:183; Franz, 1975b:223

Stigmacros froggatti (Forel) Stenichnus pusilla (King) Lea, 1912:60

Formicidae spp. Chevrolatia insignis Jacquelin Du Val Wasmann, 1894:122 0

Euconnus semiruber Casey Blatchley, 1910:294 Euconnus (Napochus) clavicorne (Casey) Blatchley, 1910:299 H Euconnus (Napochus) claviger (Muller) Markel, 1845:242

Euconnus (Napochus) fulvum (LeConte) Blatchley, 1910:300 Eutheia formicetorum Reitter Wasmann, 1894:123

Euthiconus conicicollis Reitter, 1909:222 (Fairmaire and Laboullbene)

Heterognathus carinatus King Wasmann, 1894:125



0 D

0

z

0

Table 2. Continued. ; z

Formicidae Scydmaenidae Reference 1 5

Neuraphes myrmecophilus Aube Wasmann, 1894:123 Neuraphes talparum Lokay Machulka, 1931:80 c

Scydmaenus duplicates Lea Lea, 19 10: 184 0 Scydmaenus impavidus Lea Lea, 19 12:56 Z

Scydmaenus perrisi Reitter Wasmann, 1894:125

Syndicus latipennis (Lea) Lea, 1910:188 en eI

H

0




Table 3. Scydmaenids associated with termites.

Scydmaenidae Reference

CLIDICINI Clidicus termitophilus Lhoste Lhoste, 1939:510 SCYDMAENINI Scydmaenus (Geoscydmaenus) termitophilus Franz Franz, 1971c:117 Scydmaenus (Termitoscydmaenus) kisterni Franz Franz, 1980:55 Scydmaenus biarticulatus Franz Franz, 1975d:92 Scydmaenus carinatus (King) Lea, 1910:187 Scydmaenus depigmentatus Franz Franz, 1975d: 100 Scydmaenus longipes Franz Franz, 1975d: 104 Scydmaenus sakaeratensis Franz Franz, 1975d:96 Scydmnaenus transiens Franz Franz, 1975d: 103 CYRTOSCYDMINI Euconnus butemboanus Franz Franz, 1971 c: 117 Euconnus kivuensis Franz Franz, 197 Ic: 1 17

Euconnus pilosus Cauchois Franz, 197 lc: 117 Euconnus pseudoplanocephalus Franz Franz, 1971 c: 117 Euconnus sakaeratensis Franz Franz, 1975d:79 Euconnus (Napochus) termitophilus Wasmann Wasmann, 1903:102 Stenichnus constricta (Lea) Lea, 1912:60

that Stenichnus godarti and S. collaris (Muller and Kunze) are hemimyrmecophiles (found in the vicinity of ants, but not considered as myrmecophiles), which contra dicts Donisthorpe's (1927b) observations. Neither Franc (1992) or Hlavack and Lackner (1998) provided any discussion of scydmaenid behavior or location of scyd maenids in the colony.

Most scydmaenids in the Scydmaenini, Cyrtoscydmini, and Clidicini are ant-like in appearance because they have distinct constrictions between the head and protho rax and between the prothorax and elytra, but few are as elongate and slender as

typical ants. Few have distinct morphological modifications and these are presumed to aid in copulation rather than in myrmecophily. However, the only scydmaenid that is a known myrmecophile with a specialized morphology is Plaumanniola sanc taecatharinae Costa Lima (Fig. 1), which was associated with Octostruma stenog natha Brown and Kempf (Lenko, 1972). The head is highly modified, being much

broader than long, dorso-ventrally flattened, and strongly carinate laterally, and the

terminal antennomeres are greatly enlarged, a feature typical of other known myr mecophiles (e.g., Claviger) (Holldobler and Wilson, 1990). A single specimen was collected with about 50 individuals of Octostruma Forel in leaf litter. Lenko (1972) observed that when the beetle was disturbed it curled up and remained motionless for several minutes.

Undoubtedly, most scydmaenids associated with ants are synoeketes (synoecious myrmecophiles), with few true symphiles, e.g., Euconnus chrysocomus and Plau manniola sanctaecatharinae. However, Scydmaenidae is still a poorly known group. They are undoubtedly predators, but whether they are feeding on ant eggs or larvae

or on other myrmecophiles is unknown and requires further study. Distribution of ant associates within Scydmaenidae. The family Scydmaenidae




rt

Fig. 1. Plaumanniola sanctaecatharinae Costa Lime, a myrmecophilous scydmaenid from

Brazil.




includes nearly 80 genera organized into 11 tribes. Of the 11 tribes, eight include species associated with ants. Two of the remaining tribes, Siamitini and Leptoscyd

mini, contain one and two poorly known species respectively. However, the third tribe, the Mastigini, contains about 35 species widely distributed in the Mediterra nean region and South Africa, has been well studied, and has no known ant asso ciates. Of the eight tribes with associates, the Eutheiini and Cephenniini are the most plesiomorphic, with the Chevrolatiini being somewhat more derived, but of uncertain position (Hansen, 1997). The Clidicini and Leptomastacini are closely related, with the Scydmaenini being the sister taxon to these (O'Keefe, unpubl.). The Plauman niolini is most likely a very derived form of Cyrtoscydmini, which, itself contains

many other derived groups. The Eutheiini contain five Holarctic genera, of which two, Eutheia Stephens and

Euthiconus Reitter, have associates only with European species of Formica Linnaeus, Lasius Fabricius, and Leptothorax Mayr. The Cephenniini contains ten genera that are widely distributed across the Palearctic, Nearctic, Ethiopian, and Austral-Asian regions, of which only Cephennium Muller and Kunze has three species associated

with ants, all were with Formica rufa, Lasius brunneus (Latreille), and L. fuliginosus (Latreille). The Chevrolatiini contains a single genus, Chevrolatia Jacquelin du Val, that occurs mostly in the Mediterranean region and eastern North America, of which C. insignis Jacquelin du Val has been collected with an undetermined species of ant. The Clidicini contains three poorly studied and rare scydmaenid genera. The Nearctic genus Papusus contains the only known scydmaenid associates of Pogonomyrmex

Mayr and the southeast Asian genus Clidicus contains the only associates known for

Leptogenys Roger. The Leptomastacini contains three Palearctic genera, only one of

which, the relatively well known Leptomastax Pirazzoli, has been associated with ants; they are the only scydmaenid associates known for Pheidole Westwood. The

Scydmaenini contains seven genera, of which three, Nearctic Ceramphis Casey, Aus tralian Palaeoscydmaenus Franz, and worldwide Scydmaenus, have associates with ants. Ceramphis and Palaeoscydmaenus each contain one species, and Ceramphis defornata (Horn) is the only known associate of Solenopsis Westwood (associated with S. aurea Wheeler). The genus Scydmaenus contains over 700 described species, of which 31 have been associated with ants (1 Nearctic, 4 Neotropical, 12 Palearctic, and 14 Australian) in 14 genera (Table 1). The Plaumanniolini contains a single species, the Neotropical Plaumanniola sanctaecatharinae (Fig. 1), that is the only known scydmaenid associate of Octostruma. The Cyrtoscydmini contains 46 genera (Newton and Franz, 1998), of which 10 include associates of ants. The Neotropical genera Alloraphes Franz, Mimoscydmus Franz, and Napochomorphus Franz each contains one species that is associated with Camponotus rufipes (Fabricius). The

Palearctic Neuraphes Thomson contains 15 species associated with seven genera of ants. Australian Stenichnus Thomson contains two species that are associated with six ant genera and Syndicus Motschulsky contains three species associated with ants, two of which have been associated with Iridomyrmex Mayr. Nine Palearctic species of Stenichnus and three Palearctic species of Scydmoraphes Reitter have been as

sociated with numerous ant genera. Microscydmus Saulcy and Croissandeau contains one species each in the Palearctic and Neotropics that has been associated with ants.

Euconnus Thomson contains 27 species (15 in the subgenus Napochus Thomson

alone) that have been associated with ants. Eleven of these were associated with the




Neotropical Camponotus rufipes and two others with Neotropical Odontomachus

Latreille. Three others were associated with Australian Rhytidoponera Mayr. Seven other species were associated with Palearctic ants (mostly in Formica, Lasius, and

Messor Forel) and three others were associated with Nearctic ants (all undetermined).

Ants associated with scydmaenids. Ants of four subfamilies have been associated with scydmaenid beetles: Dolichoderinae, Ponerinae, Myrmicinae, and Formicinae

(Table 2). Of the dolichoderines, only two species of Iridomyrmex and a single species of

Papyrius Shattuck have been associated with scydmaenids, and all of these were

from Australia (Clarke, 1920; Lea, 1910). Among the Ponerinae, scydmaenids, mostly in the genus Scydmaenus, have been

associated with six genera. Of these, three (Amblyopone Erichson, Pachycondyla Smith, and Rhytidoponera) are only known from work in Australia (Lea, 1910, 1912; Franz, 1975b) and one (Leptogenys) from southeast Asia associated with Clidicus (Wasmann, 1896). The only scydmaenid associate known for New World Ponerinae

are three species associated with South American Odontomachus (Franz, 1980b). Hlavac and Lackner (1998) listed the only known ponerine-scydmaenid association for the Palearctic region.

In the myrmecinae, 12 genera have been associated with scydmaenids. Six of these

(Crematogaster Lund, Leptothorax, Messor, Myrmecina Curtis, Pheidole, and Tetra

morium Mayr) are associated with Palearctic species, three (Atta Buckley, Pogono

myrmex, and Solenopsis) are associated with Nearctic scydmaenids, and one (Oc

tostruma) is associated with Neotropical scydmaenids. Only five genera in the Formicinae have been associated with scydmaenids. At

least six species of Camponotus Mayr have been associated with scydmaenids from Australia, South America, and Europe, with Camponotus rufipes (Fabricius) having been associated with 19 species of South American scydmaenids. Formica and Las ius each have numerous scydmaenid associates. Formica rufa alone has 25 associated

species of Scydmaenidae and Lasius brunneus and L. fuliginosus have 19 and 17

associated species respectively. Polyrhachis Smith and Stigmacros Forel both have

a single scydmaenid associate in Australia.

Scydmaenid-termite associations. Only 16 species of Scydmaenidae (Table 3) have been reported as being associated with termites. Eight are species of Scyd maenus, six are Euconnus, and one each in Stenichnus and Clidicus. Nearly all of

the scydmaenid-termite associations are from either Africa or Australia and were

published in either Franz' 1971 work on scydmaenids from the Congo (Franz 197 1c) or his work on southeast Asian scydmaenids (Franz 1975d). Only two Neotropical

species, Scydmaenus (Ternitoscydmaenus) kistneri Franz and Euconnus (Napochus) termitophilus Wasmann, have been associated with termites. No scydmaenid-termite associations are known from the Palearctic or Nearctic regions.

ACKNOWLEDGMENTS

I would like to thank Kevin Wiseman for drawing the figure of Plaumanniola, John Moser

for sending his notes on the locations of Scydmaenus in Atta nests, and Jerry Cook, Edward

Riley, Robert Wharton, and an anonymous reviewer for suggestions improving this paper.




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Received 15 September 1999; accepted 25 September 2000.



ant-like stone beetles, ants, and their associations (coleoptera: scydmaenidae; hymenoptera:...

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