Harpalinae

Plagiotelum irinum, from here.


Belongs within: Conjunctae.
Contains: Amarini, Odacanthini, Philophloeus, Agonocheila, Demetrida, Trigonothops, Anomotarus, Lebiini, Coptocarpus, Oodini, Chlaenius, Morion, Zuphiitae, Pterostichitae, Licinini, Harpalitae, Panagaeitae.

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).

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 [Callistitae, Coptoderinae, Lebiinae, Lebiitae, Odacanthinae, Odacanthitae]
    |--+--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
    `--+--+--+--+--Dinopelma OM08
       |  |  |  `--Hexagonia [Hexagoniinae, Hexagoniini] OM08
       |  |  |       |--H. angustula B55
       |  |  |       `--H. scabricollis B55
       |  |  `--+--Teukrus OM08
       |  |     `--Ctenodactylini LB91
       |  |          |--Leptotrachelus B90
       |  |          |--Plagiotelum Solier 1849 LM87
       |  |          |    |--*P. irinum Solier 1849 LM87
       |  |          |    `--P. opalescens Olliff 1885 LM87
       |  |          `--Ctenodactyla batesi OM08
       |  `--Zuphiitae OM08
        `--+--Pterostichitae OM08
          `--+--+--Licinini OM08
             |  `--Harpalitae OM08
             `--Panagaeitae 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 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
  Calophaena [Calophaenini] B14
    `--C. bicincta B14
         |--C. b. bicincta B14
         `--C. b. ligata Bates 1883 B14
  Calophaenoidea arrowi B14
  Graphipterus Latreille 1802 B14, L02
    |--G. circumdatus Raffr. 1885 G89
    |--G. minutus B14
    |--G. serrator (Forskål 1775) [=Carabus serrator] G89
    |--G. trilineatus [=Carabus trilineatus] L02
    `--G. variegatus [=Carabus variegatus] L02
  Ginema Ball & Shpeley 2002 [Ginemini] BS02
    `--*G. thomasi Ball & Shpeley 2002 BS02
  Amarini BS02
  Tetragonoderus Dejean 1829 BS02, LM87 [Tetragonoderinae, Tetragonoderini]
    |--*T. quadrum (Fabricius 1792) [=Carabus quadrum] LM87
    |--T. immaculatus B55
    |--T. quadrimaculatus B55
    |--T. sericatus B55
    `--T. undatus Dejean 1829 LM87
  Agra [Agriini] OM08
    |--A. aenea L02
    `--A. oblongopunctatum B90
  Calybe OM08
  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
  Clarencia Sloane 1917 LM87
    |--*C. aliena (Pascoe 1860) (see below for synonymy) LM87
    `--C. quadridens Darlington 1968 LM87
  Casnoidea Castelnau 1834 [incl. Ophionea Eschscholtz 1829 non Klug 1821] LM87
    |--*C. cyanocephala (Fabricius 1798) [=Cicindela cyanocephala] LM87
    |--‘Attelabus’ indica Thunberg 1764 [=*Ophionea indica] LM87
    |--‘Ophionea’ interstitialis B14
    |--‘Ophionea’ nigrofasciata B14
    |--C. puncticollis (Sloane 1923) [=Ophionea puncticollis] LM87
    `--C. thouzeti (Castelnau 1867) [=Ophionea thouzeti] LM87
  Renneria kamouni GWW10
  Odacanthini B12
  Somotrichus elevatus B55
  Velindopsis panagaeoides B55
  Singiliomimus modestus B55
  Neocoptodera overlaeti B55
  Parena Motschulsky 1860 [incl. Crossoglossa Chaudoir 1872, Phloeodromius Macleay 1871] LM87
    |--*P. bicolor Motschulsky 1860 LM87
    |--P. dorae Basilewsky 1955 B55
    |--P. fasciata (Chaudoir 1872) (see below for synonymy) LM87
    |--P. ferruginea Chaudoir 1878 B55
    `--P. picea (Macleay 1871) [=*Phloeodromius piceus] LM87
  Philorhizus B55
    |--P. figuratus B55
    `--P. oculatus B55
  Peliocypas B55
    |--P. debilis B55
    `--P. fragilis 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
  Stenocallida B55
    |--S. fasciata B55
    |--S. rubiginosa B55
    |--S. ruficollis B55
    `--S. umbrigera B55
  Celaenephes Schmidt-Goebel 1846 [incl. Taromorpha Blackburn 1894] LM87
    |--*C. parallelus Schmidt-Goebel 1846 LM87
    `--C. linearis (Walker 1858) [=Leistus linearis; incl. *Taromorpha alternata Blackburn 1894] LM87
  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
  Catascopus Kirby 1825 [Catascopinae] LM87
    |--*C. hardwickii Kirby 1825 LM87
    |--C. aruensis Saunders 1863 (see below for synonymy) LM87
    |--C. chalydicus Olliff 1885 LM87
    |--C. chaudoiri Castelnau 1867 LM87
    |--C. elegans (Weber 1801) (see below for synonymy) LM87
    |--C. facialis B89
    |--C. laticollis Macleay 1883 LM87
    |--C. senegalensis B55
    `--C. smaragdulus Dejean 1825 LM87
  Philophloeus 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
  Physodera Eschscholtz 1829 LM87
    |--*P. dejeani Eschscholtz 1829 LM87
    `--P. bacchusi Darlington 1971 LM87
  Lachnoderma Macleay 1873 LM87
    |--*L. cinctum Macleay 1873 LM87
    `--L. foveolatum Sloane 1915 LM87
  Demetrida LM87
  Trigonothops LM87
  Speotarus Moore 1964 LM87
    `--*S. lucifugus Moore 1964 LM87
         |--S. l. lucifugus LM87
         `--S. l. princeps Moore 1964 LM87
  Anomotarus 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
  Lebiini B14
  Cyclosomus [Cyclosomini] BS02
    |--C. flexuosus (Fabricius 1775) B14
    |--C. inustus B14
    `--C. suturalis B14
  Hololeius LaFerté-Sénectère 1851 [=Hololius Chaudoir 1857] LM87
    `--H. ceylanicus (Nietner 1856) (see below for synonymy) LM87
  Coptocarpus LM87
  Lestignathus Erichson 1842 LM87
    |--*L. cursor Erichson 1842 LM87
    |--L. foveatus Sloane 1920 LM87
    `--L. simsoni Bates 1878 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
  Oodini LM87
  Callistini LB91
  Chlaeniini [Chlaeniides, Chlaeniinae] M01
    |--Chlaenius B14
    |--Rhysotrachelus quadrimaculatus B55
    |--Ocybatus deyrollei B55
    |--Stenodinodes nepos B55
    |--Pachydinodes conformis B55
    |--Chlaenites capicola B55
    |--Pseudochlaeniellus paenulatus B55
    |--Callistoides B55
    |    |--C. hessei Basilewsky 1949 B55
    |    `--C. pulchellus B55
    |--Leptochlaenius varians B55
    |    |--L. v. varians B55
    |    |--L. v. angolanus Basilewsky 1955 B55
    |    `--L. v. verecundus B55
    |--Trachychlaenites B55
    |    |--T. gonioderus B55
    |    |    |--T. g. gonioderus B55
    |    |    `--T. g. nigrofemoralis B55
    |    `--T. obesus [incl. T. ovalipennis] B55
    |--Homalolachnus B55
    |    |--H. goossensi [incl. H. goossensi var. nigrocyanescens] B55
    |    `--H. lineatus B55
    |         |--H. l. lineatus B55
    |         `--H. l. tristis B55
    `--Chlaeniostenus B55
         |--C. denticulatus B55
         |    |--C. d. denticulatus B55
         |    `--C. d. elatus B55
         |--C. euryscopus B55
         `--C. sulcipennis B55
              |--C. s. sulcipennis B55
              `--C. s. sulcatulus B55

Catascopus aruensis Saunders 1863 [incl. C. aeneicollis Andrewes 1919, C. brevispinosus Sloane 1910, C. cupricollis Chaudoir 1883, C. dalbertisi Straneo 1943] LM87

Catascopus elegans (Weber 1801) [=Elaphrus elegans; incl. C. amoemus Chaudoir 1861, C. australasiae Hope 1842, C. elegans cyaneus Chaudoir 1872, C. obliquatus Fairmaire 1881] LM87

‘Eulebia’ picipennis Macleay 1871 [incl. Lebia papuella Darlington 1968, L. papuensis Macleay 1876] LM87

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

Parena fasciata (Chaudoir 1872) [=Crossoglossa fasciata; incl. Phloeodromius plagiatus Macleay 1876, Parena sloanei Csiki 1932] LM87

*Parophonus (Parophonus) maculicornis (Dutschmid 1812) [=Carabus maculicornis, Ophonus (*P.) maculicornis] LM87

*Type species of generic name indicated

REFERENCES

[B12] Baehr, M. 2012. A new species of the genus Pheropsophus (Coleoptera: Carabidae: Brachininae) from northern Australia. Records of the Western Australian Museum 27 (1): 62–67.

[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).

[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.

[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.

[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.

[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.

[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.

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.

Last updated: 11 June 2017.

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