Do group dynamics affect colour morph clines during a range shift?

Species exhibiting colour polymorphism are thought to have an ecological advantage at the landscape scale, because spatial segregation of alternatively adapted ecotypes into diverse habitats can increase the species' niche breadth and thus confer greater geographic range size. However, morph frequencies are also influenced by intrapopulational processes such as frequency- or density-dependent social interactions. To identify how social feedback may affect clinal variation in morph frequencies, we investigated reciprocal interactions between morph-specific thermal tolerance, local climatic conditions and social environments, in the context of a colour-morph frequency cline associated with a recent range expansion in blue-tailed damselflies (Ischnura elegans) in Sweden. Cold tolerances of gynochromes (female-like female morph) were positively correlated with local gynochrome frequencies, suggesting a positive frequency-dependent fitness benefit. In contrast, androchrome (male-mimic female morph) cold tolerances were improved following recent exposure to cold weather, suggesting a beneficial environmental acclimation effect. Thus, according to an environment-matching hypothesis for clinal variation, androchrome frequencies should therefore increase towards the (cooler) range limit. In contrast to this prediction, gynochrome frequencies increased at the expanding range limit, consistent with a positive frequency-dependent social feedback that is beneficial when invading novel climates. Our results suggest that when phenotypes or fitnesses are affected by interactions with conspecifics, beneficial social effects on environmental tolerances may (i) facilitate range shifts, and (ii) reverse or counteract typical patterns of intraspecific interactions and environment-matching clines observed in stable populations observed over broader geographic scales.