Energy density and its variation in space limit species richness of boreal forest birds

Aim An areas ability to support species may be dependent not only on the total amount of available energy it contains but also on energy density (i.e. available energy per unit area). Acknowledging these two aspects of energy availability may increase mechanistic understanding of how increased energy availability results in increased species richness. We studied the relationship between energy density, its variation in space and boreal forest bird species richness and investigated two possible mechanisms: (1) metabolic constraints of organisms, and (2) increased resource availability for specialists. Location Protected areas in Finlands boreal forest. Methods We tested whether bird species richness was best determined by total energy availability in an area or by energy density and its variation within the area, before and after including bird abundance in the models. We evaluated two main explanatory variables: tree growth reflecting the rate of energy production and tree volume as a measure of biomass. In addition, we modelled individual species responses to energy density and its variation, and evaluated the prediction of the metabolic constraints hypothesis that small species are limited by energy density whereas large species are limited by total energy availability in the area. Results Energy density and its variation were good predictors of species richness: together with abundance they explained 84% of variation in species richness (compared with 74% for abundance alone). Pure metabolic constraints were unlikely to explain this relationship. Instead, the mechanism probably involved increased habitat heterogeneity benefiting specialist species. Total energy availability was also an important factor determining species richness but its effect was indirect via abundance. Main conclusions Our results corroborate the importance of energy availability as a driver of species richness in forest bird communities, and they indicate that energy density and its variation in the landscape strongly influence species richness even after accounting for abundance.