Range shifts of overwintering birds depend on habitat type, snow conditions and habitat specialization.

Climatic warming is forcing species to shift their ranges poleward, which has been demonstrated for many taxa globally. Yet, the influence of habitat types on within- and among-species variations of distribution shifts has rarely been studied, especially during the non-breeding season. Here, we investigated habitat-specific shift distances of northern range margins and directions of the distribution center based on long-term data of overwintering birds in Finland. Specifically, we explored influences of habitat type, species' snow depth tolerance, species' climatic niche and habitat specialization on range shifts during the past 40 years in 81 bird species. Birds overwintering in arable land shifted more clearly toward north compared to birds of the same species in rural and forest habitats, while the northern range margin shift distances did not significantly differ among the habitat types. Range shifts were more linked with the species' snow depth tolerance rather than species' climatic niche. Snow depth tolerance of species was negatively associated with the eastward shift direction across all habitats, while we found habitat-specific patterns with snow depth for northward shift directions and northern margin shift distances. Species with stronger habitat specializations shifted more strongly toward north as compared to generalist species, whereas the climatic niche of bird species only marginally correlated with range shifts, so that cold-dwelling species shifted longer distances and more clearly eastward. Our study reveals habitat-specific patterns linked to snow conditions for overwintering boreal birds and highlights the importance of habitat availability and preference in climate driven range shifts.