Native Dreissena freshwater mussels in the Balkans: in and out of ancient lakes
Abstract. The Balkans is a biogeographically highly diverse region and a worldwide hotspot of endemic freshwater diversity. A substantial part of this diversity is attributed to well recognized and potential ancient lakes in its southwestern part. However, despite considerable research efforts, faunal relationships among those lakes are not well understood. Therefore, genetic information from native representatives of the mussel genus Dreissena is here used to test the biogeographical zonation of the southwestern Balkans, to relate demographic changes to environmental changes, to assess the degree of eco-insularity, to reconstruct their evolutionary history, and to explore the potential of native taxa for becoming invasive.
Phylogeographical and population genetic analyses indicate that most studied populations belong to two native species: D. presbensis (including the distinct genetic subgroup from Lake Ohrid, "D. stankovici") and D. blanci. In addition, the first confirmed record of invasive D. polymorpha in the southwestern Balkan is presented.
The distribution of native Dreissena spp. generally coincides with the biogeographical zonations previously suggested based on fish data. However, there is disagreement on the assignment of the ancient lakes in the area to respective biogeographical regions. The data for Lake Ohrid are not conclusive. A closer biogeographical connection to lakes of the Vardar region and possibly the northern Ionian region is, however, suggested for Lake Prespa.
The reconstruction of the evolutionary history of Dreissena spp. suggests that populations underwent demographic and spatial expansions in the recent past. Expansions started around 320 000–300 000 years ago in "D. stankovici", 160 000–140 000 years ago in D. blanci, and 110 000–70 000 years ago in D. presbensis. These time frames are discussed within the context of available paleogeological data for lakes Ohrid and Prespa. It is suggested that regional environmental changes may have had pronounced effects on the population histories of native Dreissena spp., though the high buffer capacity of Lake Ohrid may have lessened these effects in "D. stankovici". In addition, local events influencing individual lakes had very likely considerable effects on the demographic histories of Dreissena spp. as well. The observed patterns of immigration and emigration in and out of ancient lakes may suggest that limited gene flow enabled the survival of few isolated subpopulations and that later on eco-insularity (selective advantages of locally adopted groups) may have prevented excessive hybridization and sympatry of closely related taxa.
As for the potential invasiveness of native Dreissena spp., the inferred spatial expansions are not human-mediated and all taxa still appear to be restricted to their native ranges. A concern, however, is that today D. presbensis and D. blanci also occur in artificial water bodies, and that invasive D. polymorpha has reached the area.