Cryptic differentiation in alpine-endemic, high-altitude butterflies reveals down-slope glacial refugia

The influence of cyclic climate fluctuations and their impact on high-altitude species is still insufficiently understood. We therefore analysed in this study the genetic structure of cold-adapted animals and their coherence with geographical distributions throughout the Late Quaternary. We analysed 588 individuals from 23 populations of the alpine-endemic lesser mountain ringlet, Erebia melampus, by allozyme electrophoresis to detect its intraspecific differentiation. As an outgroup, we added one population of Erebia sudetica inalpina from Grindelwald (Swiss Alps). Seventeen of 18 loci were polymorphic. The mean F(ST) over all samples was 37%. We detected strong differentiation into three lineages with the genetic distances between the two E. melampus groups being larger than between each of the two E. melampus groups and E. sudetica. The mean genetic distance among these three groups was 0.17. These results give evidence for the existence of a species complex within the E. melampus/sudetica group and indicate a discontinuous distribution within this group during at least the last ice age. One of them, E. sudetica inalpina, is found in the northern Alps and most probably had its Würm glacial refugium north of the glaciated Alps. The western E. melampus group might have had a refugium at the southwestern Alps margin, the eastern group in the lower altitudes of the southeastern and/or eastern Alps. In the latter, a further subdivision within this relict area is possible.