Conjunctae

Nomius pygmaeus, photographed by Harvey Schmidt.


Belongs within: Geadephaga.
Contains: Cicindelinae, Paussinae, Rhysodinae, Broscinae, Amblytelus, Mecyclothorax, Harpalinae, Brachininae, Trechodini, Tasmanorites, Pogonus, Trechini, Bembidiini, Zolini.

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. The Trechinae is a group of carabid ground beetles 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 (Bell 1990).

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]
    |--+--+--Broscinae R-JC01
    |  |  `--Psydrini [Nomiini] R-JC01
    |  |       |--Nomius pygmaeus (Dejean 1831) OM08, Bo14
    |  |       `--+--Laccocenus Sloane 1890 OM08, LM87
    |  |          |    `--L. ambiguus Sloane 1890 LM87
    |  |          `--Psydrus piceus OM08, B90
    |  `--+--+--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
    |     |  `--+--Melisodera Westwood 1835 OM08, LM87 [Melisoderini R-JC01]
    |     |     |    `--*M. picipennis Westwood 1835 LM87
    |     |     `--+--Amblytelus OM08
    |     |        `--Mecyclothorax OM08
    |     `--+--Harpalinae OM08
    |        `--Brachininae OM08
    |--Trechinae [Trechitae] OM08
    |    |  i. s.: Percodermus Sloane 1920 LM87
    |    |           `--*P. niger Sloane 1920 LM87
    |    |         Sloaneana Csiki 1933 (see below for synonymy) LM87
    |    |           `--S. tasmaniae (Sloane 1915) [=Brachydema tasmaniae; incl. B. victoriae Sloane 1915] LM87
    |    |         Pterocyrtus Sloane 1920 LM87
    |    |           |--*P. globosus Sloane 1920 LM87
    |    |           |--P. rubescens Sloane 1920 LM87
    |    |           |--P. striatulus Sloane 1920 LM87
    |    |           |--P. tasmanicus (Castelnau 1867) [=Drimostoma tasmanica] LM87
    |    |           `--P. truncaticollis Sloane 1923 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
    |    |                `--P. (*P.) constrictipes (Sloane 1896) LM87 (see below for synonymy)
    |    |         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
    |    |--Pogonini [Pogoninae] R-JC01
    |    |    |  i. s.: Pachydesus OM08
    |    |    |--Octhozetus bicolor R-JC01
    |    |    `--+--Diplochaetus R-JC01
    |    |       |    |--D. planatus OM08
    |    |       |    `--D. rutilus R-JC01
    |    |       `--Pogonus R-JC01
    |    `--+--Trechini R-JC01
    |       `--+--Bembidiinae R-JC01
    |          `--Zolini R-JC01
    `--Patrobini R-JC01
         |--Patrobus OM08
         |    |--P. longicornis OM08
         |    `--P. teresae Vives & Vives 2005 F05
         |--Platidiolous vandykei B90
         |--Diplous californicus OM08
         `--Platypatrobus lacustris B90

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 [Nebriini] B14
    |--Leistus fulvibarbis (Dejean 1826) B90, VPB11
    `--Nebria Latreille 1802 L02
         |--N. brevicollis (Fabricius 1792) VPB11 [=Carabus brevicollis L02]
         |--N. gyllenhali C01
         |--N. (Boreonebria) hudsonica OM08
         |--N. livida [incl. N. livida var. lateralis, N. livida var. sibirica Csiki 1901] C01
         |--N. pallipes Say 1823 B14
         |--N. sabulosa [=Carabus sabulosus] L02
         `--N. salina Fairmaire & Laboulbene 1854 VPB11
  Tropopterus [Tropopterini] R-JC01
    |--T. giuraudyi R-JC01
    `--T. montagnii R-JC01
  Cicindelinae MW15
  0--Paussinae MW15
  `--Rhysodinae MW15

Perileptus (*Pyrrhotachys) constrictipes (Sloane 1896) LM87 [=*Pyrrotachys constrictipes LM87; incl. Pyrrotachys constricticeps Sloane 1903 LM87, Perileptus constricticeps LB91]

Sloaneana Csiki 1933 [=Brachydema Sloane 1915 non Fairmaire 1881, Sloanella Csiki 1928 non Jeannel 1927] LM87

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

*Type species of generic name indicated

REFERENCES

[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.

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

[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.

[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.

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[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.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 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.

[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.

Last updated: 23 February 2018.

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