A broader ecological context to habitat fragmentation: Why matrix habitat is more important than we thought

Consequences of habitat fragmentation have garnered much attention over the past few decades. The resulting literature has been useful for understanding how land-use changes influence population viability and community structure, but we are still hampered by a major aspect of the conceptual framework within which most fragmentation work arises. Specifically, habitat between fragments ('matrix') is usually treated as uniform and ecologically irrelevant. However, recent work on animals shows that matrix habitat can profoundly influence within-fragment dynamics. We review related evidence for plants. Various matrix types (e.g. clear-cutting, agriculture, or urbanization) can act in different ways to alter resource availability and movement of pollinators, seed dispersers, and herbivores. Inclusion of matrix qualities in fragmentation studies is further complicated since most matrices are not static; sites in which timber harvesting or agriculture occur develop through succession or change as crops are rotated, respectively, such that their influence on within-fragment processes vary temporally. Also, many plants are not restricted to remnants of original habitat. Using studies of forest understory plants, we summarize work showing how diversity can change significantly through time in matrix. Understanding the persistence of a species across fragmented landscapes will require more attention to matrix habitat, and to the species utilizing the matrix.