The spatial scales of species coexistence.

Understanding how species diversity is maintained is a foundational problem in ecology and an essential requirement for the discipline to be effective as an applied science. Ecologists' understanding of this problem has rapidly matured, but this has exposed profound uncertainty about the spatial scales required to maintain species diversity. Here we define and develop this frontier by proposing the coexistence-area relationship-a real relationship in nature that can be used to understand the determinants of the scale-dependence of diversity maintenance. The coexistence-area relationship motivates new empirical techniques for addressing important, unresolved problems about the influence of demographic stochasticity, environmental heterogeneity and dispersal on scale-dependent patterns of diversity. In so doing, this framework substantially reframes current approaches to spatial community ecology. Quantifying the spatial scales of species coexistence will permit the next important advance in our understanding of the maintenance of diversity in nature, and should improve the contribution of community ecology to biodiversity conservation.