Classification of the Genus Harpalus (Coleoptera, Carabidae) of the World Based on Imaginal Morphology

The genus Harpalus, Latreille, 1802, is the largest ground beetle genus of the tribe Harpalini and one of the most diverse genera of Carabidae (more than 400 described species). The definition and taxonomic boundaries of the genus, as well as previously proposed classifications, all based on regional faunas, are briefly reviewed in this paper from a historical perspective. The classification of the genus Harpalus proposed herein is based, for the first time, on a comparative morphological study of the world fauna and covers all the described species. The genus is divided into seventy subgenera combined into nineteen subgroups and ten groups; thirty-six subgenera are newly described: Afroharpalus subg. n. (type species Harpalus fulvicornis Thunberg, 1806), Hyloharpalus subg. n. (type species Harpalus laevipes Zetterstedt, 1828), Sinoharpalus subg. n. (type species Harpalus puetzi Kataev et Wrase, 1997), Macroharpalus subg. n. (type species Erpeinus major Motschulsky, 1850), Meroharpalus subg. n. (type species Harpalus fulvilabris Mannerheim, 1853), Ameroharpalus subg. n. (type species Harpalus spadiceus Dejean, 1829), Drymoharpalus subg. n. (type species Harpalus atratus Latreille, 1804), Caucasoharpalus subg. n. (type species Omaseus aeneipennis Faldermann, 1836), Calathoderus subg. n. (type species Harpalus potanini Tschitschérine, 1906), Isoharpalus subg. n. (type species Carabus serripes Quensel, 1806), Psammoharpalus subg. n. (type species Harpalus kozlovi Kataev), 1993, Platyharpalus subg. n. (type species Harpalus ventralis LeConte, 1848), Asioharpalus subg. n. (type species Harpalus nigrans Morawitz, 1862), Anamblystus subg. n. (type species Carabus latus Linnaeus, 1758), Homaloharpalus subg. n. (type species Carabus tardus Panzer, 1796), Bactroharpalus subg. n. (type species Harpalus cautus Dejean, 1829), Diaharpalus subg. n. (type species Harpalus vittatus Gebler, 1833), Mesoharpalus subg. n. (type species Harpalus gisellae Csiki, 1932), Eremoharpalus subg. n. (type species Harpalus remboides Solsky, 1874) Oreoharpalus subg. n. (type species Harpalus famelicus Tschitschérine, 1898), Hypsoharpalus subg. n. (type species Harpalus arnoldii Kataev, 1988), Anophonus subg. n. (type species Ophonus cyanopterus Tschitschérine, 1897), Haloharpalus subg. n. (type species Harpalus salinulus Reitter, 1900), Megaharpalus subg. n. (type species Harpalus stoetznerianus Schauberger, 1932), Aristoharpalus subg. n. (type species Harpalus ingenuus Tschitschérine, 1898), Cycloharpalus subg. n. (type species Harpalus pulvinatus Ménétries, 1848), Euryharpalus subg. n. (type species Harpalus cisteloides Motschulsky, 1844), Brachyharpalus subg. n. (type species Carabus autumnalis Duftschmid, 1812), Hemipangus subg. n. (type species Harpalus klapperichi Jedlička, 1955), Mauriharpalus subg. n. (type species Harpalus cardoni Antoine, 1922), Caloharpalus subg. n. (type species Harpalus cupreus Dejean, 1829), Baryharpalus subg. n. (type species Carabus dimidiatus Rossi, 1790), Heteroharpalus subg. n. (type species Harpalus metallinus Ménétries, 1839), Idioharpalus subg. n. (type species Harpalus numidicus Bedel, 1893), Pachyharpalus subg. n. (type species Harpalus crates Bates, 1883) and Paraharpalus subg. n. (type species Harpalus oblitus Dejean, 1829). A detailed morphological description of the genus, diagnoses, composition and distribution of all supraspecific taxa with brief biotopic notes are given.