City comfort: weaker metabolic response to changes in ambient temperature in urban red squirrels.

The ecophysiological responses of species to urbanisation reveal important information regarding the processes of successful urban colonization and biodiversity patterns in urban landscapes. Investigating these responses will also help uncover whether synurban species are indeed urban 'winners'. Yet we still lack basic knowledge about the physiological costs and overall energy budgets of most species living in urban habitats, especially for mammals. Within this context, we compared the energetic demands of Eurasian red squirrels (Sciurus vulgaris) from the core of an urban environment with those from a nearby forest. We measured oxygen consumption as a proxy for resting metabolic rate (RMR) of 20 wild individuals (13 urban, 7 forest), at naturally varying ambient temperature (T) in an outdoor-enclosure experiment. We found that the variation in RMR was best explained by the interaction between T and habitat, with a significant difference between populations. Urban squirrels showed a shallower response of metabolic rate to decreasing T than woodland squirrels. We suggest that this is likely a consequence of urban heat island effects, as well as widespread supplemental food abundance. Our results indicate energy savings for urban squirrels at cooler temperatures, yet with possible increased costs at higher temperatures compared to their woodland conspecifics. Thus, the changed patterns of metabolic regulation in urban individuals might not necessarily represent an overall advantage for urban squirrels, especially in view of increasing temperatures globally.