Phylogenetic relationships of genus Pelophila Dejean to other basal grade Carabidae (Coleoptera)

the apex, on each elytron. Otherwise, Pelophila adults could easily be mistaken for adults of some Nebria species with exceptionally short, broad, and shiny bodies and short appendages. Historically, different systematists have suggested affinities for Pelophila with Blethisa and other Elaphrini, with Nebria and other Nebriini, or, in a few cases, as an intermediate form related to both of these groups. Latreille (1802) recognized three “families” of basal carabids: the “Carabiques”, including genera now placed in the tribes Carabini and Cychrini; the “Barbus”, including Omophron, Pogonophorus (= Leistus), Loricera, and Nebria (which was described in that paper); and the “Elaphriens”, including Elaphrus and Bembidion. In 1804 (and 1810), Latreille did not consider Carabus borealis, but he placed Carabus multipunctatus Linnaeus in Nebria. Gyllenhal (1810) included both C. borealis and C. multipunctatus in Nebria. Bonelli (1810) in

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