Phylogeography of the land snail genus Circassina (Gastropoda: Hygromiidae) implies multiple Pleistocene refugia in the western Caucasus region

Abstract

The phylogeny and historical biogeography of the Caucasian land snail genus Circassina was reconstructed using multilocus amplified fragment length polymorphism (AFLP) data and mitochondrial DNA sequences. Diversification within the group started with a divergence of populations from the western Lesser Caucasus from those of the Greater Caucasus during the late Miocene. Distinct AFLP clusters and major mitochondrial clades separated by long internal branches lend evidence to the hypothesis of separate glacial refuges in the Lesser and Greater Caucasus during the Pleistocene. High genetic distances across low geographic distances and admixture analysis revealed a phylogeographic boundary running through the Colchis lowlands, which may have been established and maintained in part by repeated transgressions of the Black Sea during the Pleistocene and Holocene. Localities in Ciscaucasia were probably colonised through long-distance dispersal across the main ridge of the Greater Caucasus. The phylogeny implies multiple independent losses of accessory genital organs, i.e. dart sac and mucus glands, within Circassina. None of the anatomically defined (sub-) species distinguished so far is monophyletic and there is gene flow between the two main population groups across the Colchis lowlands. Thus, we propose to classify these population groups as subspecies of a single species.