Historical biogeography of fleas: the former Bering Land Bridge and phylogenetic dissimilarity between the Nearctic and Palearctic assemblages.

We investigated the phylogenetic structure of flea assemblages collected from small mammals on opposite sides of and increasing distance from the former Bering Land Bridge (BLB) using crossed double principal coordinate analysis (crossed-DPCoA). Phylogenetic composition of flea assemblages differed substantially between continents, but phylogenetic similarity between the Nearctic and Palearctic assemblages was the highest in the regions closer to the BLB. Within continents, phylogenetic similarity of flea assemblages was lower between regions closer to the BLB and those farther from the BLB than among regions within each of these groups. The Palearctic assemblages were represented mainly by basal families, while the Nearctic assemblages were dominated by a derived family (Ceratophyllidae), suggesting predominantly eastward pre-glaciation movements. In contrast, within the youngest flea family (Ceratophyllidae), the basal clades were characteristic for the Nearctic, while some species of a few derived clades were characteristic for the Palearctic, suggesting that, at least, during glaciation, westward movements occurred as well. In addition, multiple within-family clades of fleas were represented on opposite sides of the BLB suggesting multiple colonization events. This study is the first attempt to apply modern analytical methods of community ecology to reveal patterns in historical biogeography.