Thermal, food and vegetation effects on winter bird species richness of Mediterranean oakwoods

A better understanding of species–energy relationships needs to be developed using fine‐grained approaches that involve the use of small geographical scales of known characteristics, such as habitat heterogeneity, food availability, direct measures of temperature, and functional groups of species. We carried out a 2‐year study to analyze the effects of the thermal environment and food availability, while controlling for the influence of habitat structure, on winter species richness of birds living in oakwoods of a mountanious region of Central Spain of Mediterranean continental climate. The guild of ground‐foraging birds was selected as model organisms considering its susceptibility to winter conditions associated with unpredictable snowfalls. The spatial variation in species richness of this guild was determined by food availability, but only for those stable and predictable resources not affected by frequent snowfall (shrubs producing fruits; a complete lack of association was found with arthropod abundance on the ground). Thermal effects associated directly with air temperature, and mediated indirectly by vegetation structure providing a mosaic of sun‐shade patches, were also very influential. These patterns were highly repeatable across years. Daytime temperature had no influence on determining spatial variation in species richness, but night (minimum) temperature was a very important predictor (explained considering the lower temperatures at night, the longer duration of night, and the inability of diurnal birds to develop active behavioral thermoregulation during nighttime). This result highlights the need to consider physiological processes mediating species–environment relationships when analyzing the relationship between climatic variables and biodiversity phenomena.