Northward expanding resident species benefit from warming winters through increased foraging rates and predator vigilance.

Species distributions shift northwards due to climate change, but the ecological mechanisms allowing range expansions are not fully understood. Most studies have concentrated on breeding seasons, but winter warming may also be important. Wintering distributions are restricted by food availability and temperature, which may also interact. Foraging in cold conditions requires adaptations as individuals have to be efficient in foraging, while staying warm and vigilant for predators. When the ambient temperature declines, foraging rates should be reduced due to increased time spent on warming behaviours. In addition, predator vigilance should decline, because more time has to be invested in foraging. Cold weather should limit northward expanding southern species in particular, while northern species should perform better in cold conditions. We tested this by studying temperature responses (between 0 and - 35 °C) among wintering birds at feeders. We compared foraging behaviours of two northward expanding southern species, the great tit (Parus major) and the blue tit (Cyanistes caeruleus) to a northern species, the willow tit (Poecile montanus). Foraging rate and vigilance decreased, and warming behaviour increased when temperatures declined. Importantly, the performance in these traits was poorer in the southern species compared to the willow tit. Furthermore, the response to decreasing temperatures in foraging rates and warming behaviour was stronger in the great tits than willow tits. As the winters become warmer, these mechanisms should increase wintering success of southern species wintering at high latitudes, and lead to higher survival, increased population growth, and consequent range expansion.