Nomius pygmaeus, photographed by Harvey Schmidt.

Belongs within: Carabidae.
Contains: Nebriinae, Cicindelinae, Paussinae, Rhysodinae, Trechodini, Tasmanorites, Geocharis, Trechini, Bembidiini, Merizodinae, Pogonini, Broscinae, Brachininae, Harpalinae, Amblytelus, Mecyclothorax.

The Conjunctae are a major clade of carabid ground beetles distinguished primarily by the presence of conjunct mesocoxae, with the coxal cavities surrounded by the mesosternum and metasternum. Members of this clade include the large subfamilies Harpalinae and Trechinae, though monophyly of the latter is uncertain. A core-Trechinae clade including the Pogonini, Trechini, Bembidiini and Zolini is supported as monophyletic by the presence in males of protarsomeres 1 and 2 that are dentate and dilated on the inner side, as well as by ribosomal DNA constitution, and aschiasmatic meiosis in males (Roig-Juñent & Cicchino 2001). Other subgroups of the Conjunctae include the Patrobini, possessing a scrobal seta on the mandible and elongate outer antennal segments that are more than twice as long as wide. In contrast, Psydrini have antennae with outer segments that are oval, constricted basally and less than 1.5 times as long as wide, as well as having deep transverse groves on abdominal sterna III and IV parallel to the posterior margin (Bell 1990). Meonis is a genus of ground beetles found in Australia with elongate, porrect mandibles that may feed on snails. Melisodera, found in southeastern Australia, has moniliform antennae, globular and strongly protruding eyes, and a laterally bisetose prothorax with a distinct submarginal ridge.

See also: The diversity of ground beetles.

Synapomorphies (from Roig-Juñent & Cicchino 2001): Mesepimeron excluded from mesocoxal cavity; metepimeron forming shelf overlapping abdominal sternum II; elytral plica present; parascutellar striole absent; subapical setose organ present on gonopod; stylomere 1 or gonopod IX dimerous.

Conjunctae [Balteifera, Stylifera, Psydrinae, Psydritae, Trechinae, Trechitae]
    |--+--+--Trechini R-JC01
    |  |  `--+--Bembidiini R-JC01
    |  |     `--Merizodinae R-JC01
    |  `--Pogonini R-JC01
    |--Patrobini R-JC01
    |    |--Platidiolus Chaudoir 1878 [incl. Patroboidea Van Dyke 1925] BB01
    |    |    `--P. vandykei Kurnakov 1960 BB01
    |    |--Platypatrobus Darlington 1938 BB01
    |    |    `--P. lacustris Darlington 1938 BB01
    |    |--Diplous Motschulsky 150 BB01
    |    |    |  i. s.: D. californicus OM08
    |    |    `--D. (Platidius Chaudoir 1871) BB01
    |    |         `--D. (P.) rugicollis (Randall 1838) BB01
    |    `--Patrobus Dejean 1821 OM08, BB01
    |         |  i. s.: P. teresae Vives & Vives 2005 F05
    |         |--P. (Patrobus) [incl. Geopatrobus Darlington 1938] BB01
    |         |    |--P. (P.) fossifrons (Eschscholtz 1823) BB01
    |         |    |--P. (P.) foveocollis (Eschscholtz 1823) BB01
    |         |    `--P. (P.) septentrionis Dejean 1828 BB01
    |         `--P. (Neopatrobus Darlington 1938) BB01
    |              `--P. (N.) longicornis (Say 1823) BB01
    `--+--+--Broscinae R-JC01
       |  `--Psydrini [Nomiini] R-JC01
       |       |--Nomius Castelnau 1835 OM08, BB01 [incl. Haplochile LeConte 1848 BB01]
       |       |    `--N. pygmaeus (Dejean 1831) B14
       |       `--+--Laccocenus Sloane 1890 OM08, LM87
       |          |    `--*L. ambiguus Sloane 1890 LM87
       |          `--Psydrus LeConte 1846 OM08, BB01 [incl. Monillipatrobus Hatch 1933 BB01]
       |               `--P. piceus LeConte 1846 BB01
       `--+--+--Brachininae OM08
          |  `--Harpalinae OM08
          `--+--+--Melisodera Westwood 1835 OM08, LM87 [Melisoderini R-JC01]
             |  |    `--*M. picipennis Westwood 1835 LM87
             |  `--+--Amblytelus OM08
             |     `--Mecyclothorax OM08
             `--Meonis Castelnau 1867 OM08, LM87 [Meonidini R-JC01]
                  |--*M. niger Castelnau 1867 LM87
                  |--M. angusticollis Sloane 1911 LM87
                  |--M. ater Castelnau 1867 [incl. M. amplicollis Sloane 1915] LM87
                  |--M. convexus Sloane 1900 LM87
                  `--M. semistriatus Sloane 1916 [incl. M. minor Sloane 1916] LM87

Conjunctae incertae sedis:
  Neonomius Moore 1963 LM87
    |--*N. laevicollis (Sloane 1915) [=Mecyclothorax laevicollis] LM87
    |--N. australis (Sloane 1915) [=Mecyclothorax australis] LM87
    `--N. laticollis (Sloane 1900) [=Cyclothorax laticollis] LM87
  Sitaphe Moore 1963 LM87
    `--*S. rotundata Moore 1963 LM87
  Theprisa Moore 1963 LM87
    |--*T. convexa (Sloane 1920) [=Phersita convexa] LM87
    |--T. australis (Castelnau 1867) [=Drimostoma australis] LM87
    `--T. montana (Castelnau 1867) [=Drimostoma montana; incl. D. alpestris Castelnau 1867] LM87
  Pterogmus Sloane 1920 LM87
    `--*P. rufipes Sloane 1920 LM87
  Teraphis Castelnau 1867 [=Phersita Sloane 1903] LM87
    |--*T. melbournensis Castelnau 1867 (see below for synonymy) LM87
    |--T. cavicola Moore 1977 LM87
    |--T. crenulata (Sloane 1923) [=Phersita crenulata] LM87
    |--T. elongata Castelnau 1867 LM87
    |--T. helmsi (Sloane 1890) [=Drimostoma helmsi] LM87
    `--T. tasmanica (Sloane 1920) [=Phersita tasmanica] LM87
  Trephisa Moore 1963 LM87
    `--*T. parallela Moore 1963 LM87
  Raphetis Moore 1963 LM87
    |--*R. darlingtoni Moore 1963 LM87
    `--R. gracilis Moore 1963 LM87
  Celanida Castelnau 1867 [=Celandia (l. c.)] LM87
    `--*C. montana Castelnau 1867 LM87
  Rhaebolestes Sloane 1903 LM87
    `--*R. walkeri Sloane 1903 LM87
  Moriomorpha Castelnau 1867 LM87
    |--*M. victoriae Castelnau 1867 LM87
    `--M. adelaidae Castelnau 1867 LM87
  Moriodema Castelnau 1867 LM87
    `--*M. mcoyei Castelnau 1867 [incl. M. paramattensis Castelnau 1867] LM87
  Epelyx Blackburn 1892 LM87
    |--*E. lindensis Blackburn 1892 LM87
    `--E. latus Blackburn 1892 LM87
  Nebriinae B14
  Tropopterus [Tropopterini] R-JC01
    |--T. giuraudyi R-JC01
    `--T. montagnii R-JC01
  Cicindelinae MW15
  0--Paussinae MW15
  `--Rhysodinae MW15
  Percodermus Sloane 1920 LM87
    `--*P. niger Sloane 1920 LM87
  Trechodini B09
  Paratrechodes Jeannel 1926 LM87
    `--*P. macleayi (Sloane 1920) [=Trechus macleayi] LM87
  Trechobembix Jeannel 1926 LM87
    `--*T. baldiensis (Blackburn 1894) [=Trechus baldiensis] LM87
         |--T. b. baldiensis LM87
         |--T. b. hesperia Moore 1972 LM87
         `--T. b. queenslandica Moore 1972 LM87
  Cyphotrechodes Jeannel 1926 LM87
    `--*C. gibbipennis (Blackburn 1901) [=Trechodes gibbipennis] LM87
  Perileptus Schaum 1860 B09 [Perileptini B74]
    |  i. s.: P. africanus B55
    |         P. morimotoi Uéno 1955 I92
    |         P. sloanei Moore 1966 LM87
    |         P. westralis Moore 1972 LM87
    |--*P. (Perileptus) areolatus (Creutzer 1799) [=Carabus areolatus] LM87
    `--P. (Pyrrhotachys Sloane 1896) B09 [=Pyrrotachys LM87]
         |--P. (*P.) constrictipes (Sloane 1896) [=*Pyrrotachys constrictipes] LM87
         `--P. (P.) constricticeps (Sloane 1903) LB91, LM87 [=Pyrrotachys constricticeps LM87]
  Homoloderini B70
    |--Tasmanorites B70
    |--Pogonoschema Jeannel 1927 B70, LM87
    |    |--*P. robustum (Sloane 1920) [=Trechus robustus] LM87
    |    |--P. pallipes Moore 1972 LM87
    |    |--P. sloanei Jeannel 1927 [=Trechus pacificus Sloane 1920 non Putzeys 1870] LM87
    |    `--P. solidum Moore 1972 LM87
    `--Sloanella Jeannel 1927 B70, LM87
         |--*S. simsoni (Blackburn 1894) [=Trechus simsoni] LM87
         |--S. obscura Moore 1893 LM87
         |--S. pallida Moore 1972 LM87
         `--S. suavis Moore 1972 LM87
  Tropidotrechus Jeannel 1927 LM87
    |--*T. victoriae (Blackburn 1894) [=Trechus victoriae] LM87
    |--T. bawbawensis Moore 1972 LM87
    |--T. microps Moore 1972 LM87
    `--T. vicinus Moore 1972 LM87
  Austrotrechus Moore 1972 LM87
    |--*A. kosciuskoanus (Sloane 1923) [=Trechus kosciuskoanus] LM87
    `--A. contortus Moore 1972 LM87
  Tasmanotrechus Moore 1972 LM87
    |--*T. leai (Sloane 1920) [=Trechus leai] LM87
    |--T. cockerilli Moore 1972 LM87
    |--T. compactus Moore 1983 LM87
    `--T. concolor Moore 1972 LM87
  Geodetrechus Moore 1972 LM87
    |--*G. talpinus Moore 1972 LM87
    |--G. mendumae Moore 1972 LM87
    `--G. parallelus Moore 1972 LM87
  Nototrechus Moore 1972 LM87
    `--*N. unicolor Moore 1972 LM87
  Trechistus Moore 1972 LM87
    |--*T. humicola Moore 1972 LM87
    |--T. inconspicuus Moore 1972 LM87
    |--T. stenoderus Moore 1972 LM87
    |--T. sylvaticus Moore 1972 LM87
    `--T. terricola Moore 1972 LM87
  Trechiella Jeannel 1927 LM87
    |--*T. subornatella (Blackburn 1901) [=Trechus subornatellus] LM87
    `--T. queenslandica Moore 1972 LM87
  Mimanillus Moore 1972 LM87
    `--*M. gracilis Moore 1972 LM87
  Eutrechopsis Moore 1972 LM87
    `--*E. ovalis Moore 1972 LM87
  Mimotrechus Moore 1972 LM87
    |--*M. australiensis (Sloane 1923) [=Trechus australiensis] LM87
    |--M. carteri (Sloane 1920) [=Trechus carteri] LM87
    |--M. obscuroguttatus Moore 1972 LM87
    `--M. scitulus Moore 1972 LM87
  Bothynotrechus Moore 1972 LM87
    |--*B. castelnaui (Sloane 1920) [=Trechus castelnaui] LM87
    `--B. lynx Moore 1972 LM87
  Geocharis Z05

*Teraphis melbournensis Castelnau 1867 [=*Phersita melbournensis; incl. T. argutoroides Castelnau 1867] LM87

*Type species of generic name indicated


[B09] Baehr, M. 2009. A new species of the tachyine genus Tasmanitachoides from the Kimberley Division, Western Australia (Coleoptera, Carabidae). Records of the Western Australian Museum 25 (2): 159–164.

[BB01] Ball, G. E., & Y. Bousquet. 2001. Carabidae Latreille, 1810. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 32–132. CRC Press: Boca Raton.

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

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[B70] Britton, E. B. 1970. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 495–621. Melbourne University Press.

[B74] Britton, E. B. 1974. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 62–89. Melbourne University Press.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

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

[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 543–683. Melbourne University Press: Carlton (Victoria).

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.

[OM08] Ober, K. A., & D. R. Maddison. 2008. Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8: 63.

[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.

[Z05] Zaballos, J. P. 2005. Los Geocharis Ehlers, 1883 de Marruecos y Cádiz (España) (Coleoptera, Carabidae, Trechinae, Anillini). Graellsia 61 (1): 61–81.

Last updated: 22 December 2020.

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