The geometry of habitat fragmentation: effects of distribution patterns on short-term species persistence
Land-use changes cause habitat loss and fragmentation and are thus important drivers of anthropogenic biodiversity change. However, there is an ongoing debate about how fragmentation per se affects biodiversity in a given amount of habitat. We illustrate why it is important to distinguish two different aspects of fragmentation to resolve this debate: (i) geometric fragmentation effects, which exclusively arise from the spatial distributions of species and habitat fragments, and (ii) demographic fragmentation effects due to reduced fragment size, increased isolation, or edge effects. While most empirical studies are primarily interested in quantifying demographic fragmentation effects, geometric effects are typically invoked only as post-hoc explanations of biodiversity responses to fragmentation per se. Here, we present an approach to quantify geometric fragmentation effects on species persistence probability. We illustrate this approach using spatial simulations where we systematically varied the initial abundances and distribution patterns (i.e. random, aggregated, and regular) of species as well as habitat amount and fragmentation per se. As expected, we found no geometric fragmentation effects when species were randomly distributed. However, when species were aggregated, we found positive effects of fragmentation per se on persistence probability for a large range of scenarios. For regular species distributions, we found weakly negative geometric effects. These findings are independent of the ecological mechanisms which generate non-random species distributions. Our study helps to reconcile seemingly contradictory results of previous fragmentation studies. Since intraspecific aggregation is a ubiquitous pattern in nature, our findings imply widespread positive geometric fragmentation effects. This expectation is supported by many studies that find positive effects of fragmentation per se on species occurrences and diversity after controlling for habitat amount. We outline how to disentangle geometric and demographic effects of fragmentation, which is critical for predicting the response of biodiversity to landscape change.