The integripennis species group of Geocharidius Jeannel, 1963 (Carabidae, Bembidiini, Anillina) from Nuclear Central America: a taxonomic review with notes about biogeography and speciation

Our review recognizes 15 species of the integripennis species group of Geocharidius from Nuclear Central America, include three species previously described (Geocharidiusgimlii Erwin, Geocharidiusintegripennis (Bates) and Geocharidiuszullinii Vigna Taglianti) and 12 described here as new. They are: Geocharidiusandersoni sp. n. (type locality: Chiapas, Chiapas Highlands, Cerro Huitepec) and Geocharidiusvignatagliantii sp. n. (type locality: Chiapas, Motozintla, Sierra Madre de Chiapas, Benito Juárez) from Mexico; Geocharidiusantigua sp. n. (type locality: Sacatepéquez, 5 km SE of Antigua), Geocharidiusbalini sp. n. (type locality: Suchitepéquez, 4 km S of Volcan Atitlán), Geocharidiuserwini sp. n. (type locality: Quiché Department, 7 km NE of Los Encuentros), Geocharidiusjalapensis sp. n. (type locality: Jalapa Department, 4 km E of Mataquescuintla), Geocharidiuslonginoi, sp. n. (type locality: El Progreso Department, Cerro Pinalón), and Geocharidiusminimus sp. n. (type locality: Sacatepéquez Department, 5 km SE of Antigua) from Guatemala; and Geocharidiuscelaquensis sp. n. (type locality: Lempira Department, Celaque National Park), Geocharidiuscomayaguanus sp. n. (type locality: Comayagua Department, 18 km ENE of Comayagua), Geocharidiusdisjunctus sp. n. (type locality: Francisco Morazán, La Tigra National Park), and Geocharidiuslencanus sp. n. (type locality: Lempira Department, Celaque National Park) from Honduras. For all members of the group, adult structural characters, including male and female genitalia, are described, and a taxonomic key for all members of the integripennis species group is presented based on these characters. Behavioral and biogeographical aspects of speciation in the group are discussed, based on the morphological analysis. In all cases of sympatry, pairs of closely related species show greater differences in sizes than pairs of more remotely related species. Integripennis group species occupy six different montane areas at elevations above 1300m, with no species shared among them. Major faunal barriers in the region limiting present species distributions include the Motagua Fault Zone and a gap between the Guatemalan Cordillera volcanic chain and the Honduran Interior Highlands no higher than 900m in elevation. Highest species diversity is in the Guatematan Cordillera (six species), second highest in the Honduran Interior Highlands area (four species).