The species-area-energy relationship.

Area and available energy are major determinants of species richness. Although scale dependency of the relationship between energy availability and species richness (the species-energy relationship) has been documented, the exact relationship between the species-area and the species-energy relationship has not been studied explicitly. Here we show, using two extensive data sets on avian distributions in different biogeographic regions, that there is a negative interaction between energy availability and area in their effect on species richness. The slope of the species-area relationship is lower in areas with higher levels of available energy, and the slope of the species-energy relationship is lower for larger areas. This three-dimensional species-area-energy relationship can be understood in terms of probabilistic processes affecting the proportions of sites occupied by individual species. According to this theory, high environmental energy elevates species' occupancies, which depress the slope of the species-area curve.