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Key Words habitat loss, landscape scale, habitat configuration, patch size, patch isolation, extinction threshold, landscape complementation
Abstract The literature on effects of habitat fragmentation on biodiversity is huge. It is also very diverse, with different authors measuring fragmentation in different ways and, as a consequence, drawing different conclusions regarding both the magnitude and direction of its effects. Habitat fragmentation is usually defined as a landscape-scale process involving both habitat loss and the breaking apart of habitat. Results of empirical studies of habitat fragmentation are often difficult to interpret because (a) many researchers measure fragmentation at the patch scale, not the landscape scale and (b) most researchers measure fragmentation in ways that do not distinguish between habitat loss and habitat fragmentation per se, i.e., the breaking apart of habitat after controlling for habitat loss. Empirical studies to date suggest that habitat loss has large, consistently negative effects on biodiversity. Habitat fragmentation per se has much weaker effects on biodiversity that are at least as likely to be positive as negative. Therefore, to correctly interpret the influence of habitat fragmentation on biodiversity, the effects of these two components of fragmentation must be measured independently. More studies of the independent effects of habitat loss and fragmentation per se are needed to determine the factors that lead to positive versus negative effects of fragmentation per se. I suggest that the term "fragmentation" should be reserved for the breaking apart of habitat, independent of habitat loss.
INTRODUCTION
A recent search of the Cambridge Scientific Abstracts database revealed over 1600 articles containing the phrase "habitat fragmentation." The task of reviewing this literature is daunting not only because of its size but also because different authors use different definitions of habitat fragmentation, and they measure fragmentation in different ways and at different spatial scales.
This diversity of definitions of habitat fragmentation can be readily seen in the titles of some articles. For example, "Impacts of habitat fragmentation and patch size..." (Collingham & Huntly 2000) suggests that habitat fragmentation and patch size are two different things. However, other authors actually use patch size to measure habitat fragmentation (e.g., Golden & Crist 2000, Hovel & Lipicus 2001). "The effects of forest fragmentation and isolation..." (Goodman & Rakotodravony 2000) suggests that forest fragmentation...