Geadephaga

Scarites eurytus, photographed by A. Anichtchenko.

Belongs within: Adephaga.
Contains: Pterostichini, Cicindelinae, Conjunctae, Pseudomorphinae, Zuphiinae.

The Geadephaga is a lineage of terrestrial, mostly predatory, adephagan beetles supported, albeit weakly, by molecular analysis. Most Geadephaga are classified in the large and diverse family Carabidae, but the group also includes the small relictual family Trachypachidae, whose members lack the rocking hind coxae of the Carabidae (Grimaldi & Engel 2005).

<==Geadephaga [Caraboidea]
    |--Trachypachidae GE05
    |    |--Unda cursoria P02
    |    |--Karatoma agilis P02
    |    |--Evertus Z02
    |    |--Systolosoma brevis R-JC01
    |    |--Trachypachus GE05
    |    `--Psacodromeus guttata GE05
    `--Carabidae R-JC01
         |  i. s.: Omophron M01
         |         Omus D03
         |         Metrius D03
         |         Carabilarva R02
         |         Protorabus planus P02
         |         Chlaenius M01
         |           |  i. s.: C. planicornis B66
         |           |         C. spoliatus G89
         |           |         C. tenellus G89
         |           `--C. (Diapheropsophus) B66
         |                |--C. (D.) mellyi B66
         |                `--C. (D.) swinhoei Bates 1866 B66
         |         Nebria VPB11
         |           |--N. brevicollis (Fabricius 1792) VPB11
         |           `--N. salina Fairmaire & Laboulbene 1854 VPB11
         |         Ctenognathus novaezealandiae Fairmaire 1842 H82
         |         Clivina B89
         |           |--C. parryi Putzeys 1863 [incl. Eupalamus clivinoides] B89
         |           `--C. rugithorax M83
         |         Euthenaris M83
         |         Notagonum submetallicum M83
         |         Scopodes elaphroides M83
         |         Pelecium A71
         |         Megadromus E02
         |           |--M. antarcticus (Chaudoir 1865) E02
         |           |--M. rectangulus WH02
         |           `--M. temukensis WH02
         |         Holcaspis WH02
         |           |--H. augustula WH02
         |           |--H. delator WH02
         |           `--H. subaenea WH02
         |         Creobius MC94
         |         Cascellius MC94
         |         Triplosarus E04
         |         Carabus [Carabina, Carabini] E04
         |           |  i. s.: C. intricatus Linnaeus 1761 VPB11
         |           |         C. nemoralis E04
         |           |         C. violaceus CM07
         |           `--C. (Ohomopterus) SS06
         |                |--C. (O.) dehaanii SS06
         |                |--C. (O.) iwawakianus SS06
         |                |--C. (O.) uenoi (Ishikawa 1960) SS06, I92
         |                |--C. (O.) yaconinus SS06
         |                `--C. (O.) yamato SS06
         |         Dromius E04
         |         Cychrus AM07
         |         Procerus AM07
         |         Calosoma BHM08
         |         Notiophilus GE05
         |           |--N. biguttatus (Fabricius 1779) VPB11
         |           `--N. quadripunctatus (Dejean 1826) VPB11
         |         Triaenogenius G89
         |           |--T. anthioides (Chaud. 1872) [=Acanthogenius anthioides] G89
         |           |--T. arabicus Gestro 1889 G89
         |           |--T. carinulatus (Fairm. 1887) [=Acanthogenius carinulatus] G89
         |           |--T. corpulentus Chaud. 1877 G89
         |           |--T. ferox (Erichs. 1843) [=Helluo ferox] G89
         |           |--T. helluonoides (Aney 1881) [=Anthia helluonoides] G89
         |           `--T. sculpturatus (Gerst. 1866) [=Acanthogenius sculpturatus] G89
         |         Pheropsophus africanus [=Brachinus africanus] G89
         |         Brachinus B89
         |           |--B. caligatus Bates 1889 B89
         |           |--B. frontalis B89
         |           `--B. oblongus G89
         |         Glycia castanea [=Cymindis castanea] G89
         |         Trichis pallida G89
         |         Blechrus G89
         |           |--B. infuscatus Motsch 1859 [incl. B. stricticollis Baudi 1864] G89
         |           `--B. plagiatus [=Lebia plagiata] G89
         |         Lyonichus G89
         |           |--L. beccarii Gestro 1889 [=Lionychus beccarii] G89
         |           `--L. sulcatus Chaud. 1876 G89
         |         Siagona G89
         |           |--S. brunnipes G89
         |           `--S. europaea G89
         |         Graphipterus G89
         |           |--G. circumdatus Raffr. 1885 G89
         |           `--G. serrator (Forsk. 1775) [=Carabus serrator] G89
         |         Distichus planus (Bonelli 1813) [=Scarites planus] G89
         |         Eccoptomenus eximius [=Chlaenius eximius] G89
         |         Penthimus obscurus G89
         |         Crasodactylus punctatus G89
         |         Dichirotrichus obsoletus [=Harpalus (Ophonus) obsoletus] G89
         |         Stenolophus B89
         |           |--S. cyanellus Bates 1889 B89
         |           |--S. gonidius Bates 1889 B89
         |           |--S. marginatus B89
         |           `--S. quinquepustulatus B89
         |         Abacetus B89
         |           |--A. aeneus [=Feronia (Argutor) aenea] G89
         |           |--A. amplicollis Bates 1889 B89
         |           |--A. bisignatus Bates 1889 B89
         |           |--A. quadriguttatus B89
         |           |--A. quadripustulatus B89
         |           `--A. salzmanni B89
         |         Poecilus conformis G89
         |         Orthomus longulus (Reiche 1855) [=Feronia longula] G89
         |         Syrdenus grayi (Woll. 1862) [=Pogonus grayi; incl. P. fulvus Baudi 1864] G89
         |         Epicosmus B89
         |           |--E. feae Bates 1889 B89
         |           `--E. transversalis B89
         |         Lamprophonus Bates 1889 B89
         |           `--*L. lucens Bates 1889 B89
         |         Notiobia B89
         |         Liodaptus Bates 1889 B89
         |           `--*L. birmanicus Bates 1889 B89
         |         Agonoderus B89
         |         Daptus B89
         |         Oxycentrus Chaud. 1854 [incl. Aegreuter Schmidt-Goeb. 1846 (n. n.)] B89
         |           |--O. angustus Bates 1876 [incl. Argreuter melas Schmidt-Goeb. 1846 (n. n.)] B89
         |           `--O. parallelus B89
         |         Anoplogenius B89
         |           |--A. microgonus B89
         |           `--A. rutilans Bates 1889 B89
         |         Perigona ruficollis Motsch 1851 B89
         |         Trigonotoma B89
         |           |--T. concinna B89
         |           |--T. dohrnii B89
         |           `--T. lewisii [incl. T. l. var. bhamoensis Bates 1889] B89
         |         Triplogenius B89
         |           |--T. chalcothorax B89
         |           `--T. mouhotii B89
         |         Drimostoma rectangulum Chaudoir 1872 B89
         |         Morio orientalis B89
         |         Loxandrus B89
         |           |--L. birmanus Bates 1889 B89
         |           `--L. brevicollis B89
         |         Pirantillus Bates 1889 B89
         |           `--*P. feae Bates 1889 B89
         |         Cyphocoleus B89
         |         Onycholabis B89
         |         Glyptolenus B89
         |         Drypta B89
         |           |--D. aeneipennis Bates 1889 B89
         |           |--D. distincta B89
         |           `--D. fulveola Bates 1883 I92
         |         Orthogonius B89
         |           |--O. davidis B89
         |           |--O. duplicatus [incl. O. alternans] B89
         |           `--O. profundestriatus B89
         |         Masoreus B89
         |           |--M. adelioides [=Aephnidius adelioides] B89
         |           |--M. fuscipennis [=Aephnidius fuscipennis] B89
         |           `--M. simplex [=Aephnidius simplex] B89
         |         Peripristus ater [=Thyreopterus ater] B89
         |         Catascopus facialis B89
         |         Dolichoctis B89
         |           |--D. ornatellus B89
         |           `--D. rotundatus B89
         |         Coptodera elegantula B89
         |         Anchista binotata (see below for synonymy) B89
         |         Plagiotelum AY04
         |         Pamborus AY04
         |         Maoripamborus AY04
         |         Loricera pilicornis (Fabricius 1775) VPB11
         |         Tachyta N88
         |         Schizogenius N88
         |         Strategus N88
         |         Calathus N88
         |         Pterostichini L88
         |         Antilliscaris L88
         |         Prodyscherus L88
         |         Agonum L88
         |           |--A. decorum L88
         |           |--A. dorsalis VPB11
         |           |--A. elongatulum L88
         |           `--A. moestum (Dufts 1812) VPB11
         |         Phloeoxena L88
         |         Idacarabus G60
         |         Badister bipustulatus (Fabricius 1792) VPB11
         |         Asaphidion VPB11
         |           |--A. flavipes (Linnaeus 1761) VPB11
         |           `--A. stierlini (Heyden 1880) VPB11
         |         Diachromus germanus (Linnaeus 1758) VPB11
         |         Leistus fulvibarbis (Dejean 1826) VPB11
         |         Oodes helopioides (Fabricius 1792) VPB11
         |         Metromenus O94
         |--Cicindelinae GE05
         |--Conjunctae R-JC01
         |--Rhysodinae GE05
         |--Pseudomorphinae B08
         |--Zuphiinae B86
         |--Paussus A61 [Paussinae GE05]
         |    `--P. lineatus A61
         |--Lebia A71 [Lebiinae GE05]
         |    `--L. scapularis A71
         |--Anchomeninae B89
         |    |--Megalonychus B89
         |    |--Euleptus B89
         |    `--Feanus Bates 1889 B89
         |         `--*F. spinipennis Bates 1889 B89
         `--Scaritinae R-JC01
              |--Migadops [Migadopini] R-JC01
              |    `--M. latus R-JC01
              `--Scarites G89
                   |--S. eurytus G89
                   |--S. gigas C74
                   `--S. guineensis [incl. S. guineensis var. asphaltinus] G89

Anchista binotata [=Plochionus binotatus; incl. Calleida discophora Chaudoir 1852, Paraphoea signifera Bates 1873] B89

*Type species of generic name indicated

REFERENCES

[AM07] Acosta, L. E., & G. Machado. 2007. Diet and foraging. In Harvestmen: The Biology of Opiliones (R. Pinto-da-Rocha, G. Machado & G. Giribet, eds) pp. 309-338. Harvard University Press: Cambridge (Massachusetts).

[A61] Andreae, H. 1961. Coleoptera: Cossyphodidae. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 198-216. Almqvist & Wiksell: Uppsala.

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales & G. S. Taylor. 2004. Insects 'Down Under' - diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216-234.

[B86] Baehr, M. 1986. Revision of the Australian Zuphiinae. 6. The genus Planetes Macleay. Supplement to the other genera (Insecta, Coleoptera, Carabidae). Spixiana 9 (2): 151-168.

[B08] Baehr, M. 2008. New and rare pseudomorphine species from Western Australia (Insecta, Coleoptera, Carabidae, Pseudomorphinae). Records of the Western Australian Museum 24 (2): 151-180.

[BHM08] Baert, L., F. Hendrickx & J.-P. Maelfait. 2008. Allozyme characterization of Hogna species (Araneae, Lycosidae) of the Galápagos Archipelago. Journal of Arachnology 36 (2): 411-417.

[B66] Bates, H. W. 1866. On a collection of Coleoptera from Formosa sent home by R. Swinhoe, Esq., H. B. M. Consul, Formosa. Proceedings of the Zoological Society of London 1866: 339-355.

[B89] Bates, H. W. 1889. Viaggio di Leonardo Fea in Birmani e regioni vicine. XVI. – On some Carabidae from Burma collected by Mr. L. Fea. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 100-111.

[C74] Coineau, Y. 1974. Éléments pour une monographie morphologique, écologique et biologique des Caeculidae (Acariens). Mémoires du Muséum National d'Histoire Naturelle, nouvelle série, Série A, Zoologie 81: 1-299, 224 pls.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In Harvestmen: The Biology of Opiliones (R. Pinto-da-Rocha, G. Machado & G. Giribet, eds) pp. 339-373. Harvard University Press: Cambridge (Massachusetts).

[D03] Didham, R. K. 2003. New invertebrate phylogenies. Weta 25: 1-7.

[E02] Emberson, R. M. 2002. The beetle (Coleoptera) fauna of the Chatham Islands: Additions and corrections. New Zealand Entomologist 25: 69-77.

[E04] Emberson, R. M. 2004. A European ground beetle, Harpalus tardus (Panzer) (Coleoptera: Carabidae), in Canterbury, and the predisposition of European carabids to establish in exotic temperate locations. New Zealand Entomologist 27: 39-42.

[G60] Gatenby, J. B. 1960. The Australasian mycetophilid glowworms. Transactions of the Royal Society of New Zealand 88 (3): 577-593.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880. – IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a 7: 5-72.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H82] Helmore, D. W. 1982. Drawings of New Zealand insects. Bulletin of the Entomological Society of New Zealand 8: 1-52.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[L88] Liebherr, J. K. 1988. General patterns in West Indian insects, and graphical biogeographic analysis of some circum-Caribbean Platynus beetles (Carabidae). Systematic Zoology 37 (4): 385-409.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: An annotated species list. DSIR Entomology Division Report 3: 1-98.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45-92.

[MC94] Morrone, J. J., & J. M. Carpenter. 1994. In search of a method for cladistic biogeography: An empirical comparison of component analysis, Brooks parsimony analysis, and three-area statements. Cladistics 10: 99-153.

[N88] Noonan, G. R. 1988. Biogeography of North American and Mexican insects, and a critique of vicariance biogeography. Systematic Zoology 37 (4): 366-384.

[O94] Otte, D. 1994. The Crickets of Hawaii: origin, systematics and evolution. The Orthopterists' Society: The Academy of Natural Sciences of Philadelphia.

[P02] Ponomarenko, A. G. 2002. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 164-176. Kluwer Academic Publishers: Dordrecht.

[R02] Rasnitsyn, A. P. 2002. Scope and approach. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 1-8. Kluwer Academic Publishers: Dordrecht.

[R-JC01] Roig-Juñent, S., & A. C. Cicchino. 2001. Chaltenia patagonica, new genus and species belonging to Chalteniina, a new subtribe of Zolini (Coleoptera: Carabidae). Canadian Entomologist 133: 651-670.

[SS06] Sota, T. & M. Sasabe. 2006. Utility of nuclear allele networks for the analysis of closely related species in the genus Carabus, subgenus Ohomopterus. Systematic Biology 55 (2): 329-344.

[VPB11] Varet, M., J. Pétillon & F. Burel. 2011. Comparative responses of spider and carabid beetle assemblages along an urban-rural boundary gradient. Journal of Arachnology 39 (2): 236-243.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.