How the spatial scales of dispersal, competition, and environmental heterogeneity interact to affect coexistence

Spatial coexistence depends on a variety of biological and physical processes, and the relative scales of these processes may promote or suppress coexistence. We model plant competition in a spatially varying environment to show how shifting scales of dispersal, competition, and environmental heterogeneity affect coexistence. Spatial coexistence mechanisms are partitioned into three types: the storage effect, nonlinear competitive variance, and growth-density covariance. We first describe how the strength of each of these mechanisms depends on covariances between population densities and between population densities and the environment, and we then explain how changes in the scales of dispersal, competition, and environmental heterogeneity should affect these covariances. Our quantitative approach allows us to show how changes in the scales of biological and physical processes can shift the relative importance of different classes of spatial coexistence mechanisms and gives us a more complete understanding of how environmental heterogeneity can enable coexistence. For example, we show how environmental heterogeneity can promote coexistence even when competing species have identical responses to the environment.