ISSN 0013-8738, Entomological Review, 2013, Vol. 93, No. 3, pp. 370–397. © Pleiades Publishing, Inc., 2013. Original Russian Text © B.M. Kataev, 2012, published in Entomologicheskoe Obozrenie, 2012, Vol. 91, No. 4, pp. 800–831. 370 Ground-Beetles of the Subgenus Cryptophonus Brandm. et Z. Brandm., Genus Harpalus Latr. (Coleoptera, Carabidae) B. M. Kataev Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia e-mail:harpal@zin.ru Received August 20, 2012 Abstract–A taxonomic review of ten species of the subgenus Cryptophonus Brandmayr et Zetto Brandmayr, 1982, the genus Harpalus Latreille, 1802 is given. In addition to nine Palaearctic species, the Ethiopian H. agnatus Rei-che, 1849 is also included in Cryptophonus. Harpalus cyrenaicus Koch, 1939, stat. n. from Libya, which was origi-nally described as a subspecies of H. litigiosus Dejean, 1829, is treated as a distinct species. Harpalus tenebrosus Dejean, 1829 is treated as a polytypical species with two subspecies: H. t. tenebrosus (West Palaearctic) and H. t. paivanus Wollaston, 1867, stat. n. (Cabo Verde). A new substitute name H. melancholicus reicheianus nom. n. is proposed to replace the objective homonyms H. reichei Jacobson, 1907 (non Desbrochers des Loges, 1867) and H. ovalis Reiche, 1861 (non Motschulsky, 1844). Lectotypes are designated for H. tenebrosus Dejean, 1829, H. paivanus Wollaston, 1867, H. litigiosus Dejean, 1829, H. agnatus Reiche, 1849, H. fulvus Dejean, 1829, H. melancholicus Dejean, 1829, and H. ineditus Dejean, 1829. The status and diagnosis of Cryptophonus are dis-cussed and a key to all the species of this subgenus is provided. Data on distribution of each species are provided. The following species are recorded for the first time: H. tenebrosus from Afghanistan, H. grilli Kataev, 2002 from India (Uttarhand), H. agnatus Reiche, 1849 from Tanzania, and H. fulvus Dejean, 1829 from Portugal. DOI: 10.1134/S001387381303010X Cryptophonus Brandmayr et Zetto Brandmayr, 1982 has been established as a subgenus of the genus Ophonus Dejean, 1821 for four West Palaearctic spe-cies of Harpalus Latreille, 1802 (H. tenebrosus De-jean, 1829, H. litigiosus Dejean, 1829, H. melan-cholicus Dejean, 1829, and H. fulvus Dejean, 1829) despite their typical “harpaloid” appearance and ab-sence of the pubescence on dorsum typical for Ophonus. Such treatment was based on the “opho-noid”-type of larval features of H. tenebrosus (larvae of three other species are unknown), in particular of presence of two pronounced teeth on anterior margin of clypeus characteristic of Ophonus (Brandmayr et al., 1980; Brandmayr and Zetto Brandmayr, 1982). In addition, some peculiar imaginal features of these species, first of all the presence of reduced pubescence on the dorsal surface of tarsi, were also taken into account. In the former classifications (Jeannel, 1942; Antoine, 1959), all the mentioned species were treated as a natural group of uncertain rank within Harpalus based mainly on their aedeagi lacking distinct apical capitulum. Sciaky (1987) in his revision of the West Palaearctic Ophonus treated Cryptophonus as a sepa-rate genus because this taxon, in his opinion, distinctly differs from Ophonus in characters of both external morphology and the structure of male genitalia, but unfortunately he did not indicate these differences. Sciaky (l. c.) added three Atlantic island species (Har-palus schaumii Wollaston, 1864, H. janinae Jeanne, 1984 and H. paivanus Wollaston, 1867) to Crypto-phonus and suggested that this genus should also in-clude some more Asian and African species described within Harpalus. Kataev (in Kryzhanovskij et al., 1995) indicated some additional distinctive characters for Cryptophonus and ranked it as a species group within the genus Harpalus; the genus Ophonus was treated by him as distinct from Harpalus on the basis of imaginal morphology. More recently (1997, 2002) he included two Himalayan species (H. idiotus Bates, 1889 and H. grilli Kataev, 2002) into Cryptophonus. Based on the morphological, caryological and mito-chondrial cytochrome-oxidase data, Martíinez-Navarro et al. (2003, 2005) considered Cryptophonus as a sepa-rate taxon well differentiated either from Harpalus and Ophonus. However, this opinion is based on data derived from a very restricted number of species and,
