High molecular diversity in Carabus (Hygrocarabus) variolosus and C. nodulosus

Keywords: diversity, COI, ITS2, species versus subspecies, introgression, refuges, Carabus (Hygrocarabus) variolosus, Carabus nodulosus


The Carabus subgenus Hygrocarabus contains two taxa: C. variolosus and C. nodulosus, the species or subspecies status of which is handled far from uniform in the literature. Both taxa show a similar morphology, the shape of the tip of the aedeagus provides a reliable morphological marker for identification. We analysed two mitochondrial gene parts (COI-5’ and COI-3’) and a nuclear one (ITS2). High diversity was found showing specific geographical patterns. Introgressive hybridisation was detected but interpreted not as an argument for subspecies status because high genetic distances indicated that it must have taken place in former times. In a laboratory hybridisation experiment, the male did not accept the female of the other taxon, supporting the conclusion that these are separate species. A series of refuges was expected for the period of ice ages. Although only the taxon C. variolosus is listed in Annex II and IV of the EU Habitats Directive, C. nodulosus also falls under this listing, as at the time of including the species into the Annexes in 2004, the two taxa were considered subspecies and hence the listing would include both, independent of later taxonomic revisions.