The story of endurance: Biogeography and the evolutionary history of four Holarctic butterflies with different habitat requirements

Aim Biogeographical studies on the entire ranges of widely distributed species can change our perception of species' range dynamics. We studied the effects of Pleistocene glacial cycles on current butterfly species distributions, aiming to uncover complex biogeographic patterns in the Holarctic, a region dramatically affected by Cenozoic climate change. Location Eurasia and North America. Taxon Boloria chariclea, Agriades optilete, Carterocephalus palaemon, Oeneis jutta. Methods We reconstructed the biogeographic history of four butterfly species differing in habitat preferences (B. chariclea - tundra, A. optilete - bogs, C. palaemon - temperate grasslands, O. jutta - taiga), using one mitochondrial and two nuclear DNA markers and species distribution modelling. Results Except for B. chariclea, all species originated in Eurasia. The open habitat species A. optilete and C. palaemon formed widely distributed east-west genetic clusters in continental Asia and clusters separated from them in Europe. Genetic clusters of the taiga species O. jutta were not geographically separated in Eurasia, suggesting Pleistocene fragmentation and recent reconnection. The glaciated North America was recolonized from Beringian and southerly situated refugia by all four species. Main conclusions The Pleistocene mammoth steppe allowed a widespread continuous distribution of open habitat butterflies, while in contrast the distribution of a taiga-specialist species was more limited. In the mostly flat and continental North Asia, the butterflies of various types of open habitats survived ice age in widely distributed east-west belts. In the mountainous and oceanic regions of Europe, Beringia and west North America, all four species persisted in contracted areas during the glacials. After deglaciation, they expanded their ranges and formed contact zones among populations. To conclude, the harsh climate of the glacials did not represent an obstacle for butterflies. Instead, different habitat specialists selected their own ways to thrive in the dynamic conditions of Quaternary glacial periods.