Occupancy Is Nine-Tenths of the Law: Occupancy Rates Determine the Homogenizing and Differentiating Effects of Exotic Species

Biotic homogenization, the loss of local biotic distinctiveness among locations (beta diversity), is a form of global change that can result from the widespread introduction of nonnative species. Here, we model this process using only species’ occupancy rates—the proportion of sites they occupy—without reference to their spatial arrangement. The nonspatial model unifies many empirical results and reliably explains >90% of the variance in species’ effects on beta diversity. It also provides new intuitions and principles, including the conditions under which species’ appearance, spread, or extirpation will homogenize or differentiate landscapes. Specifically, the addition or spread of exotic species that are more common than the native background rate (effective occupancy) homogenizes landscapes, while driving such species to extinction regionally or introducing rarer species differentiates them. Given the primacy of occupancy and our model’s ability to explain its role, homogenization research can now focus on other factors.