Theory meets reality: How habitat fragmentation research has transcended island biogeographic theory

Island biogeography theory (IBT) provides a basic conceptual model for understanding habitat fragmentation. Empirical studies of fragmented landscapes often reveal strong effects of fragment area and isolation on species richness, although other predictions of the theory, such as accelerated species turnover in fragments, have been tested less frequently. As predicted by IBT, biota in fragments typically 'relax' over time towards lower species richness. Beyond these broad generalizations, however, the relevance of IBT for understanding fragmented ecosystems is limited. First, IBT provides few predictions about how community composition in fragments should change over time, and which species should be most vulnerable. Second, edge effects can be an important driver of local species extinctions and ecosystem change, but are not considered by IBT. Third, the matrix of modified vegetation surrounding fragments-also ignored by IBT-can strongly influence fragment connectivity, which in turn affects the demography, genetics, and survival of local populations. Fourth, most fragmented landscapes are also altered by other anthropogenic changes, such as hunting, logging, fires, and pollution, which can interact synergistically with habitat fragmentation. Finally, fragmentation often has diverse impacts on ecosystem properties such as canopy-gap dynamics, carbon storage, and the trophic structure of communities that are not considered by IBT. I highlight these phenomena with findings from fragmented ecosystems around the world. Published by Elsevier Ltd.