High mountain ranges such as the European Alps and Pyrenees were the last to deglaciate in mainland Europe during glacial minima over the last ~700 ky (Darnault et al., 2012;Ehlers et al., 2018).For taxa adapted to warm climates, the persistence and extent of Alpine ice sheets and the east-west orientation of the mountain ranges hindered recolonization from more southerly refugia in Italy and the Iberian Peninsula into Europe (Taberlet et al., 1998). Coldtolerant organisms exhibit a more complex history. Many expanded their range during glacial periods, some persisted in nunataks (e.g., ice-free rocky protrusions) within the Alps, and trans-Alpine recolonization has occurred in several species (e.g., Oak species, the common shrew, and barred grass snake, and the European stag beetle;