Next, we aimed to identify those climatic factors, which directly relate to genetic divergence and thus possibly contribute to divergence or cohesion of lineages across Europe. The association of the 19 bioclimatic variables to genetic differentiation was evaluated using GESTE (Foll and Gaggiotti 2006). In addition to the bioclimatic variables, we included longitude and latitude for all populations in the analysis, to test for isolation by distance. GESTE was run with 10 pilot runs and a sample size of 10 000, with a thinning interval of 20 and a burnin of 50 000.