Geadephaga

Trachypachus inermis, copyright University of Alaska Museum.


Belongs within: Adephaga.
Contains: Geocharis, Scaritinae, Anchomeninae, Elaphrinae, Conjunctae, Carabinae.

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; a small restricted group whose members lack the rocking hind coxae of typical Carabidae have commonly been separated as a distinct family Trachypachidae but McKenna et al.'s (2015) phylogenetic analysis nested them within the Carabidae. Modern trachypachids are restricted to the Holarctic Trachypachus and South American Systolosoma (Grimaldi & Engel 2005). The Mesozoic Eodromeinae have mostly been included in the Trachypachidae but a phylogenetic analysis by Beutel et al. (2013) placed at least some eodromeines outside the geadephagan crown clade. A similar position was occupied by the Jurassic aquatic genus Timarchopsis, previously classified in the extinct dytiscoid family Coptoclavidae. Classification within the Carabidae has historically been confused with authors differing substantially in their preferred arrangements.

<==Geadephaga [Geodephaga]
    |--Eodromeinae BW13
    |    |--Sogdodromeus LM87
    |    |--Karadromeus BW13
    |    |--Eodromeus robustus BW13
    |    |--Sinodromeus liutiaogouensis Wang, Zhang & Ponomarenko 2012 BW13
    |    `--Ademosynoides asiaticus Martynov 1936 BW13
    `--+--Timarchopsis Brauer, Redtenbacher & Ganglbauer 1989 [=Necronectes Ponamarenko 1977 (preoc.)] BW13
       |    `--T. aquaticus (Ponomarenko 1977) BW13
       `--Carabidae [Carabici, Cicindeletae, Cicindelidae, Simplicia] BW13
            |  i. s.: Carabilarva R02
            |         Protorabus planus P02
            |         Ctenognathus novaezealandiae Fairmaire 1842 H82
            |         Holcaspis WH02
            |           |--H. augustula WH02
            |           |--H. delator WH02
            |           `--H. subaenea WH02
            |         Creobius MC94
            |         Cascellius MC94
            |         Triplosarus E04
            |         Procerus AM07
            |         Glycia castanea [=Cymindis castanea] G89
            |         Trichis pallida G89
            |         Blechrus C01
            |           |--B. infuscatus Motsch 1859 [incl. B. stricticollis Baudi 1864] G89
            |           `--B. minutulus C01
            |         Lyonichus G89
            |           |--L. beccarii Gestro 1889 [=Lionychus beccarii] G89
            |           `--L. sulcatus Chaud. 1876 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
            |         Orthomus longulus (Reiche 1855) [=Feronia longula] G89
            |         Syrdenus grayi (Woll. 1862) [=Pogonus grayi; incl. P. fulvus Baudi 1864] G89
            |         Lamprophonus Bates 1889 B89
            |           `--*L. lucens Bates 1889 B89
            |         Liodaptus Bates 1889 B89
            |           `--*L. birmanicus Bates 1889 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
            |         Trigonotoma [Trigonotomides] B89
            |           |--T. concinna B89
            |           |--T. dohrnii B89
            |           `--T. lewisii [incl. T. l. var. bhamoensis Bates 1889] B89
            |         Drimostoma rectangulum Chaudoir 1872 B89
            |         Pirantillus Bates 1889 B89
            |           `--*P. feae Bates 1889 B89
            |         Cyphocoleus B89
            |         Onycholabis B89
            |         Glyptolenus B89
            |         Peripristus ater [=Thyreopterus ater] B89
            |         Anchista binotata (see below for synonymy) B89
            |         Maoripamborus AY04
            |         Antilliscaris L88
            |         Prodyscherus L88
            |         Diachromus germanus (Linnaeus 1758) VPB11
            |         Metromenus O94
            |         Geocharis SA04
            |         Apoduvalius (Apoduvalius) anseriformis Salgado & Peláez 2004 F05
            |         Platyderus F05
            |           |--P. marduk Anichtchenko 2004 F05
            |           `--P. preciosae Campos & Novoa 2005 F05
            |         Typhlocharis F05
            |           |--T. fozcoaensis Serrano & Aguiar 2005 F05
            |           |--T. gonzaloi Ortuño 2005 F05
            |           `--T. passosi Serrano & Aguiar 2005 F05
            |         Eobroscus lutshniki (Roubal 1928) L98
            |         Promecognathus [Promecognathini] B90
            |         Gehringia [Gehringiini] B90
            |           `--G. olympica B90
            |         Opisthius [Opisthiini] B90
            |           `--O. richardsoni B90
            |         Lachnophorini B90
            |           |--Ega B90
            |           |--Euphorticus pubescens B90
            |           `--Lachnophorus B90
            |         Inna [Eucheilini] B90
            |           `--I. texana B90
            |         Thlibops [Scapterini] B55
            |         Lorocera pilicornis C01
            |         Tachypus C01
            |           |--T. flavipes C01
            |           `--T. pallipes C01
            |         Pseudotaphoxenus dauricus C01
            |         Zabrus trinii C01
            |         Ophonus Stephens 1827 LM87
            |           |--O. azureus [incl. O. azureus var. coeruleipennis] C01
            |           |--O. calceatus C01
            |           |--O. cordatus C01
            |           |--O. griseus C01
            |           `--O. pubescens C01
            |         Corsyra fusula C01
            |         Lionedrya mongolica C01
            |         Brachynus C01
            |           |--B. barbarus Lucas 1846 E12
            |           |--B. explodens [incl. B. explodens var. glabratus] C01
            |           |--B. fimbriolatus Lucas 1846 E12
            |           |--B. immaculicornis C01
            |           `--B. sclopeta C01
            |         Pseudophonus ruficornis (Fabricius 1775) [=Carabus ruficornis, Harpalus ruficornis] LM87
            |         Fortiseode pervalimand Jia et al. 2011 BW13
            |         Scaritinae R-JC01
            |         Anchomeninae B55
            |         Elaphrinae B14
            |         Loricera Latreille 1802 [Loricerinae, Loricerini] L02
            |           `--*L. pilicornis (Fabricius 1775) L02, VPB11 [=Carabus pilicornis L02]
            |         Thyreopterinae B55
            |           |--Thyreopterus flavosignatus [incl. T. flavosignatus var. lyratus] B55
            |           `--Thyreopterinus overlaeti [incl. T. overlaeti var. vaneyeni] B55
            |         Cicindinae B14
            |           |--Cicindis horni Bruch 1908 B14
            |           `--Archaeocindis johnbeckeri B14
            |         Collyris B66 [Collyridinae G07]
            |           |--C. filiformis Chaud. 1864 B66
            |           |--C. formosana Bates 1866 B66
            |           `--C. loochooensis Kano 1929 TYM08
            |         Siagona [Siagoninae] B14
            |           |--S. brunnipes G89
            |           |--S. dejeani B14
            |           `--S. europaea Dejean 1826 B14
            |         Omophroninae [Omophronini] B55
            |           |--Phrator vittulatus B55
            |           `--Omophron B55
            |                |--O. congoense B55
            |                |--O. limbatum [=Scolytus limbatus] L02
            |                `--O. tesselatum Say 1823 B14
            |         Orthogonius [Orthogoniinae] B55
            |           |--O. brevilabris B55
            |           |--O. clarkei B55
            |           |--O. davidis B89
            |           |--O. duplicatus [incl. O. alternans] B89
            |           `--O. profundestriatus B89
            |         Hiletinae B14
            |           |--Hiletus B14
            |           `--Eucamaragnathus B14
            |                |--E. amapa B14
            |                |--E. batesi (Chaudoir 1861) B14
            |                |--E. brasiliensis B14
            |                `--E. jaws B14
            |         Graphopterus [Graphopterinae] B55
            |           |--G. circumcinctus B55
            |           |    |--G. c. circumcinctus B55
            |           |    `--G. c. lineolatus B55
            |           |--G. congoensis B55
            |           |    |--G. c. congoensis B55
            |           |    `--G. c. monardi B55
            |           `--G. laticollis B55
            |         Pogonophorus Latreille 1802 L02
            |           `--*P. spinibarbis [=Carabus spinibarbis] L02
            |--Conjunctae MW15
            `--+--Carabinae MW15
               `--Trachypachini [Trachypachidae, Trachypachinae] MW15
                    |--Unda cursoria P02
                    |--Karatoma agilis P02
                    |--Evertus Z02
                    |--Systolosoma brevis R-JC01
                    |--Trachypachus inermis Motschulsky 1850 B14
                    `--Psacodromeus guttata GE05

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

*Type species of generic name indicated

REFERENCES

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[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.

[B55] Basilewsky, P. 1955. Carabidae (Coleoptera) de l’Angola (première partie). Diamang Publicações Culturais 27: 91–137.

[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.

[B90] Bell, R. T. 1990. Insecta: Coleoptera Carabidae adults and larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1053–1092. John Wiley & Sones: New York.

[BW13] Beutel, R. G., B. Wang, J.-J. Tan, S.-Q. Ge, D. Ren & X.-K. Yang. 2013. On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera, Insecta). Cladistics 29 (2): 147–165.

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[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

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[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 2a, 7: 5–72.

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[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.

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[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

[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: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.

Last updated: 11 June 2017.

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