Harpalinae

Badister peltatus, copyright Wim Rubers.


Belongs within: Conjunctae.
Contains: Agonocheila, Trigonothops, Coptocarpus, Morion, Zuphiitae, Pterostichitae, Licinini, Harpalitae, Zabrini, Odacanthini, Pseudomorphini, Platynini, Oodini, Perigona, Lebiini.

The Harpalinae is a large clade of carabid ground beetles, distinguished by the distinctive forms of the parameres in the male genitalia. Over 19,000 species of Harpalinae are known from the modern fauna, exhibiting a wide range of ecologies including herbivorous and myrmecophilous forms. These species have been arranged in a correspondingly large number of tribes, but relationships between the tribes have been poorly known (Ober & Maddison 2008). These include the Morionini, a group of species with moniliform antennae, a projecting dentiform process above the antennal insertion, and usually protruding eyes.

Synapomorphies (from Roig-Juñent & Cicchino 2001): Mandibular scrobal setae absent; maxillary palpomere 4 fusiform; elytral plica absent; left paramere conchiferous, differing in shape from right; right paramere glabrous.

<==Harpalinae (see below for synonymy)
    |--+--Moriosomus OM08
    |  `--Morionini [Morionides] B14
    |       |--Morion OM08
    |       |--Megamorio B14
    |       |--Hyperion Castelnau 1834 B14, LM87 (see below for synonymy)
    |       |    `--*H. schroetteri (Schreibers 1802) (see below for synonymy) LM87
    |       `--Platynodes westermanni Westwood 1847 B14
    |            |--P. w. westermanni B14
    |            `--P. w. peregrinus B14
    `--+--+--+--Hexagoniini [Hexagoniinae] OM08
       |  |  |    |--Dinopelma immaculatum OM08
       |  |  |    `--Hexagonia OM08
       |  |  |         |--H. angustula B55
       |  |  |         `--H. scabricollis B55
       |  |  `--Ctenodactylini OM08
       |  |       |--Teukrus cruciatus OM08
       |  |       `--+--Leptotrachelus dorsalis OM08
       |  |          `--Ctenodactyla batesi OM08
       |  `--Zuphiitae OM08
        `--+--Pterostichitae OM08
          `--+--+--Licinini OM08
             |  `--Harpalitae OM08
             `--+--Zabrini OM08
                `--+--+--Peleciini [Peleciinae] OM08
                   |  |    |--Pelecium sulcipenne OM08
                   |  |    |--Disphericus tarsalis B55
                   |  |    `--Eripus nitidus OM08
                   |  `--Odacanthini OM08
                   `--+--+--Pseudomorphini OM08
                      |  `--Platynini OM08
                      `--+--+--+--Calophaena [Calophaenini] OM08
                         |  |  |    |--C. bicincta B14
                         |  |  |    |    |--C. b. bicincta B14
                         |  |  |    |    `--C. b. ligata Bates 1883 B14
                         |  |  |    `--C. dupuisi OM08
                         |  |  `--Lachnophorini OM08
                         |  |       |  i. s.: Ega B90
                         |  |       |--Aporesthus anomalus OM08
                         |  |       `--+--Asklepia OM08
                         |  |          `--+--+--Anchonoderus OM08
                         |  |             |  `--Euphorticus pubescens OM08, B90
                         |  |             `--+--Calybe laetula OM08
                         |  |                `--Lachnophorus elegantulus OM08
                         |  `--Oodini OM08
                         `--+--+--+--Sinurus opacus OM08
                            |  |  `--Stenotelus opacus OM08
                            |  `--+--+--Cyrtolaus L88
                            |     |  `--+--Barylaus L88
                            |     |     `--Caelostomus Macleay 1825 OM08, LM87 (see below for synonymy)
                            |     |          |--*C. picipes (Macleay 1825) (see below for synonymy) LM87
                            |     |          |--C. albertisi Straneo 1938 LM87
                            |     |          `--C. subsinuatus (Chaudoir 1883) [=Drimostoma subsinuata] LM87
                            |     `--+--Badister Clairville 1806 OM08, LM87
                            |        |    |  i. s.: B. neopulchellus OM08
                            |        |    |         B. reflexus OM08
                            |        |    |--*B. (Badister) bipustulatus (Fabricius 1792) LM87 (see below for synonymy)
                            |        |    `--B. (Baudia Ragusa 1884) LM87
                            |        |         |--B. (*B.) peltatus (Panzer 1796) [=Carabus peltatus] LM87
                            |        |         `--B. (B.) sundaicus Andrewes 1926 LM87
                            |        `--Perigonini [Perigoninae, Perigonitae] OM08
                            |             |--Perigona OM08
                            |             `--+--Diploharpus laevissimus OM08
                            |                `--Ripogena bembidioides [=Perigona (Ripogena) bembidioides] OM08
                            `--Lebiini OM08

Harpalinae incertae sedis:
  Cillenum Samouelle 1819 B27
    |--C. chalmeri B27
    `--C. tillyardi Brookes 1927 B27
  Dichaetochilus Basilewsky 1946 B55
    |--D. incrassatus B55
    |--D. moestus B55
    `--D. planicollis B55
  Hyparpalus B55
    |--H. conviva B55
    `--H. holosericeus B55
  Laparhetes gracilis B55
  Platymetopus B55
    |--P. figuratus B55
    `--P. platythorax B55
  Neosiopelus Basilewsky 1946 B55
    `--N. punctatellus B55
  Aulacoryssus B55
    |--A. natalicus B55
    `--A. venustulus B55
  Xenodochus melanarius B55
    |--X. m. melanarius B55
    `--X. m. sissekabanus Basilewsky 1946 B55
  Ooidius dorsiger B55
  Bradybaenus Dejean 1829 B55, BR17 [Bradybaenina]
    `--B. opulentus B55
         |--B. o. opulentus B55
         `--B. o. insularis B55
  Microlestia B55
    |--M. atrata B55
    |--M. cribricollis B55
    `--M. immerita B55
  Netrodera formicaria B55
    |--N. f. formicaria B55
    |--N. f. damarensis B55
    `--N. f. peringueyi B55
  Thermophilum Basilewsky 1950 B55
    |--T. crudele B55
    |    |--T. c. crudele B55
    |    |--T. c. bennigseni B55
    |    `--T. c. westermanni B55
    |--T. homoplatum B55
    `--T. limbatum B55
         |--T. l. limbatum B55
         |--T. l. machadoi Basilewsky 1955 B55
         `--T. l. ovampoa B55
  Brachygnathus B14
    |--B. angusticollis Burmeister 1885 B14
    `--B. oxygonus B14
  Calophaenoidea arrowi B14
  Ginema Ball & Shpeley 2002 [Ginemini] BS02
    `--*G. thomasi Ball & Shpeley 2002 BS02
  Gestroania Liebke 1938 LM87
    `--*G. amplipennis (Gestro 1875) [=Casnonia amplipennis] LM87
  Myrmecodemus Sloane 1923 LM87
    |--*M. riverinae (Sloane 1890) [=Casnonia riverinae] LM87
    |--M. formicoides (Sloane 1910) [=Lachnothorax formicoides] LM87
    `--M. globulicollis (Macleay 1888) [=Casnonia globulicollis] LM87
  Anasis Castelnau 1867 LM87
    `--*A. howittii Castelnau 1867 LM87
  Aulacolius Sloane 1923 LM87
    `--*A. triordinatus Sloane 1923 LM87
  Deipyrus Liebke 1938 LM87
    `--*D. palustris (Sloane 1910) [=Lachnothorax palustris] LM87
  Basistichus Sloane 1917 LM87
    `--*B. micans (Macleay 1864) [=Odacantha micans] LM87
  Renneria kamouni GWW10
  Velindopsis panagaeoides B55
  Singiliomimus modestus B55
  Neocoptodera overlaeti B55
  Philorhizus B55
    |--P. figuratus B55
    `--P. oculatus B55
  Coptoderina B55
    |--C. amoenula B55
    |--C. congoana B55
    `--C. rugiceps B55
  Metalebia B55
    |--M. antelobata Basilewsky 1955 B55
    |--M. promontorii B55
    |--M. transvaalensis B55
    `--M. vaciva B55
  Coptoglossus Chaudoir 1869 LM87
    |--*C. sulcatulus Chaudoir 1869 LM87
    `--C. carteri (Sloane 1915) [=Platynus carteri] LM87
  Holcoderus Chaudoir 1869 LM87
    |--*H. praemorsus Chaudoir 1869 LM87
    `--H. caeruleipennis Sloane 1910 LM87
  Agonocheila LM87
  Minuthodes Andrewes 1941 [=Platia Chaudoir 1869 non Hübner 1820] LM87
    |--*M. lineella (Chaudoir 1869) [=*Platia lineella] LM87
    `--M. queenslandica (Sloane 1917) [=Platia queenslandica] LM87
  Mochtherus Schmidt-Goebel 1846 LM87
    `--M. tetraspilotus (Macleay 1825) (see below for synonymy) LM87
  Dolichoctis Schmidt-Goebel 1846 LM87
    |--*D. striata Schmidt-Goebel 1846 [incl. Mochtherus macleayi Sloane 1907] LM87
    |--D. aculeata Chaudoir 1869 LM87
    |--D. ornatellus B89
    |--D. rotundatus B89
    `--D. subrotunda Darlington 1968 LM87
  Trigonothops LM87
  Speotarus Moore 1964 LM87
    `--*S. lucifugus Moore 1964 LM87
         |--S. l. lucifugus LM87
         `--S. l. princeps Moore 1964 LM87
  Aristolebia Bates 1892 LM87
    |--*A. quadridentata Bates 1892 LM87
    `--A. wau Darlington 1968 LM87
  Creagris Nietne 1857 [incl. Pseudhelluo Castelnau 1867] LM87
    |--*C. labrosa Nietner 1857 LM87
    `--C. wilsonii (Castelnau 1867) [=*Pseudhelluo wilsonii] LM87
  Helluonidius Chaudoir 1872 LM87
    |--*H. cyanipennis (Hope 1842) (see below for synonymy) LM87
    |--H. aterrimus (Macleay 1873) [=Helluosoma aterrimum] LM87
    |--H. cyaneus (Castelnau 1867) [=Helluosoma cyanea] LM87
    `--H. latipennis (Macleay 1887) [=Helluosoma latipenne] LM87
  Helluosoma Castelnau 1867 (see below for synonymy) LM87
    |--*H. atrum Castelnau 1867 [=H. ater] LM87
    `--H. longicolle Macleay 1888 LM87
  Helluodema Castelnau 1867 [incl. Simoglossus Chaudoir 1872] LM87
    |--H. unicolor (Hope 1842) (see below for synonymy) LM87
    `--H. brunneum Sloane 1917 LM87
  Epimicodema Sloane 1914 LM87
    `--*E. mastersii (Macleay 1871) [=Helluosoma mastersii] LM87
  Helluapterus Sloane 1914 LM87
    `--*H. niger Sloane 1914 [incl. H. niger punctata Sloane 1914] LM87
  Neohelluo Sloane 1914 LM87
    `--*N. angulicollis Sloane 1914 LM87
  Ametroglossus Sloane 1914 LM87
    `--*A. ater (Macleay 1887) [=Gigadema atrum] LM87
  Hololeius LaFerté-Sénectère 1851 [=Hololius Chaudoir 1857] LM87
    `--H. ceylanicus (Nietner 1856) (see below for synonymy) LM87
  Coptocarpus LM87
  Siagonyx Macleay 1871 LM87
    |--*S. amplipennis Macleay 1871 LM87
    |--S. blackburni Sloane 1916 LM87
    `--S. mastersii Macleay 1871 LM87
  Hormacrus Sloane 1898 LM87
    |--*H. latus Sloane 1898 LM87
    `--H. minor (Blackburn 1890) [=Lestignathus minor] LM87
  Platylytron Macleay 1873 [=Platelytron Csiki 1931] LM87
    `--*P. amplipenne Macleay 1873 LM87
  Dilonchus Andrewes 1936 LM87
    |--*D. bidens Andrewes 1936 LM87
    `--D. pictus Darlington 1963 LM87
  Omestes Andrewes 1933 LM87
    `--*O. torta Andrewes 1933 LM87
  Callistini LB91

*Badister (Badister) bipustulatus (Fabricius 1792) LM87 [=Carabus bipustulatus LM87, Licinus bipustulatus L02]

Caelostomus Macleay 1825 OM08, LM87 [incl. Stomonaxus Motschulsky 1860 LM87; Caelostomini]

*Caelostomus picipes (Macleay 1825) [=Anaulacus (*Caelostomus) picipes; incl. Drimostoma rufipes Boheman 1858, Caelostomus striaticollis Chaudoir 1872] LM87

Harpalinae [Callistitae, Coptoderinae, Lebiinae, Lebiitae, Licinina, Masoreidae, Masoreomorphi, Odacanthinae, Odacanthitae]

Helluodema unicolor (Hope 1842) [=Aenigma unicolor; incl. Helluomorpha batesii Thomson 1857, *Helluodema batesii, Gigadema castelnaui Chaudoir 1872, *Simoglossus niger Chaudoir 1872] LM87

*Helluonidius cyanipennis (Hope 1842) [=Aenigma cyanipenne; incl. Helluosoma viridipenne Macleay 1887] LM87

Helluosoma Castelnau 1867 [incl. Helluodema Chaudoir 1872 non Castelnau 1867, Helluodoma (l. c.)] LM87

Hyperion Castelnau 1834 B14, LM87 [=Campylocnemis Westwood 1842 LM87, Heteroscelis Boisduval 1835 non Latreille 1829 LM87]

*Hyperion schroetteri (Schreibers 1802) [=Scarites schroetteri, *Campylocnemis schroetteri, *Heteroscelis schroetteri] LM87

Mochtherus tetraspilotus (Macleay 1825) [=Dromius tetraspilotus; incl. *M. angulatus Schmidt-Goebel 1846] LM87

*Type species of generic name indicated

REFERENCES

[BS02] Ball, G. E., & D. Shpeley. 2002. Ginemini, Ginema thomasi, new tribe, new genus and new species, from Amazonian Bolivia (Coleoptera: Carabidae: Harpalinae). Transactions of the American Entomological Society 128 (1): 75–98.

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

[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.

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

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[B27] Brookes, A. E. 1927. A new genus and three new species of Coleoptera. Transactions and Proceedings of the New Zealand Institute 57: 563–566.

[GWW10] Guthrie, N. A., T. Weir & K. Will. 2010. Localised and regional patterns in ground-dwelling beetle assemblages in a semi-tropical arid zone environment. Records of the Western Australian Museum Supplement 78 (1): 169–184.

[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.

[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.

[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.

Last updated: 7 March 2018.

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