Phylogenetic relationships and evolution of the Japanese carabinae ground beetles based on mitochondrial ND5 gene sequences

The phylogenetic relationships of the Japanese Carabinae ground beetles were analyzed by comparing 1,069 nucleotide sequences in the mitochondrial gene encoding NADH dehydrogenase subunit 5 (ND5). The ND5 phylogenetic tree revealed that the hind-wingless Carabina and the hind-winged Calosoma/Campalita (Calosomina) diverged from the common ancestor, and Cychrus (Cychrini) is the outgroup of them. Five distinct clusters (groups) can be recognized in the Carabina, i.e., CARABUS, HEMICARABUS, LEPTOCARABUS, APOTOMOPTERUS, and PROCRUSTES/DAMASTER. The ancestors of these lineages diverged almost at the same time more than 10 Myr ago. The Carabus cluster includes two subclusters, Carabus and Ohomopterus. Two species of Carabus examined are phylogenetically rather remote, while five species among Ohomopterus are closely related to each other. The results suggest that diversification of Carabus started much earlier than that of Ohomopterus, presumably in the Eurasian continent, and that of Ohomopterus in the Japanese archipelago. The branching order in the LEPTOCARABUS lineage was established, Authenocarabus/Pentacarabus being their outgroup. In the DAMASTER/PROCRUSTES lineage, Procrustes is placed as the outgroup of Damaster, with the branching order of Coptolabrus and Acoptolabrus/Damaster. The diversification of the Damaster subspecies appeared to have occurred in the Japanese archipelago earlier than Ohomopterus, and its phylogeny reflects their geographic distribution in the archipelago rather than the morphological characters.