Climate change pushes species polewards and upwards – as temperatures rise, species move to areas that were previously too cold for them. During range expansions, species encounter unfamiliar environmental conditions, which may require evolutionary adaptation, but expanding populations may often be hampered by their genetic and demographic properties. Whether range-expanding populations can adapt may greatly affect species distributions, but the question is largely unexplored for native species expanding in response to climate change. 

In seasonal environments, organisms must endure harsh conditions and synchronise growth and reproduction with the presence of food and mates. To time their life cycles, many animals and plants use seasonal changes in daylength. Insects typically overwinter in diapause (dormancy with paused development and suppressed metabolism), which is induced by short days well before winter. But across-latitude differences in daylength pose challenges for latitudinal range expansions. I focus on whether traits related to seasonal timing and winter survival have evolved during range expansion of the wall brown butterfly (Lasiommata megera) in Sweden.

In Chapter I, I confirmed that the wall brown has, in 2000–2020, expanded northwards along the eastern and western coasts of Sweden, and in Chapter II, I demonstrated that these parallel expansions have proceeded independently from the south, in isolation from each other. Laboratory experiments in Chapter I revealed that caterpillars from northern populations have evolved to correctly interpret their local daylength cues. This rapid evolution, repeated along two range expansions, indicates that latitudinal differences in daylength may seldom hinder insect range expansions. In Chapter II, I found that northern range margin populations have lower genetic variation than southern ones but are unlikely to have received much locally maladaptive gene flow from the south. Further, a genomic scan suggested that the parallel phenotypic changes have evolved through non-parallel genetic changes. In Chapter III, a laboratory experiment showed lack of local adaptation to different winters, in contrast to the rapid evolution of diapause timing. Overall winter survival was low in the coldest treatment, indicating that winter temperatures limit the range.

In Chapter IV, I studied diapause induction, growth rate, and winter survival in a field setting. Almost all individuals entered diapause, with only minimal impact from the among-population differences found in Chapter I. These evolved differences could stem from natural selection on earlier parts of the generation, which experience longer days than our experiment captured. Further, individuals of northern descent grew faster than those from the south. This could help them grow large enough before winter, yet pre-winter mass did not affect winter survival. This time, natural selection may favour high growth rates in late-hatching individuals with less time to grow before winter. Like in Chapter III, there was no evidence for evolution of improved winter survival, and survival dropped markedly when transplanting individuals outside of the current range.

Despite rapid evolution in two traits, cold winters limit the wall brown’s expansion. To predict range expansions, we must pinpoint their drivers, study trait evolution relevant to these drivers, and recognize that traits that are crucial in different seasons may vary in evolvability.