Structure of the species--energy relationship.

The relationship between energy availability and species richness (the species-energy relationship) is one of the best documented macroecological phenomena. However, the structure of species distribution along the gradient, the proximate driver of the relationship, is poorly known. Here, using data on the distribution of birds in southern Africa, for which species richness increases linearly with energy availability, we provide an explicit determination of this structure. We show that most species exhibit increasing occupancy towards more productive regions (occurring in more grid cells within a productivity class). However, average reporting rates per species within occupied grid cells, a correlate of local density, do not show a similar increase. The mean range of used energy levels and the mean geographical range size of species in southern Africa decreases along the energy gradient, as most species are present at high productivity levels but only some can extend their ranges towards lower levels. Species turnover among grid cells consequently decreases towards high energy levels. In summary, these patterns support the hypothesis that higher productivity leads to more species by increasing the probability of occurrence of resources that enable the persistence of viable populations, without necessarily affecting local population densities.