Dispersal increases ecological selection by increasing effective community size

Selection and drift are universally accepted as the cornerstones of evolutionary changes. Recent theories extend this view to ecological changes, arguing that any change in species composition is driven by deterministic fitness differences among species (enhancing selection) and/or stochasticity in birth and death rates of individuals within species (enhancing drift). These forces have contrasting effects on the predictability of ecological dynamics, and thus understanding the factors affecting their relative importance is crucial for understanding ecological dynamics. Here we test the hypothesis that dispersal increases the relative importance of ecological selection by increasing the effective size of the community (i.e., the size relevant for competitive interactions and drift). According to our hypothesis, dispersal increases the effective size of the community by mixing individuals from different localities. This effect diminishes the relative importance of demographic stochasticity, thereby reducing drift and increasing the relative importance of selective forces as drivers of species composition. We tested our hypothesis, which we term the "effective community size" hypothesis, using two independent experiments focusing on annual plants: a field experiment in which we manipulated the magnitude of dispersal and a mesocosm experiment in which we directly manipulated the effective size of the communities. Both experiments, as well as related model simulations, were consistent with the hypothesis that increasing dispersal increases the role of selective forces as drivers of species composition. This finding has important implications for our understanding of the fundamental forces affecting community dynamics, as well as the management of species diversity, particularly in patchy and fragmented environments.