Toward multifactorial null models of range contraction in terrestrial vertebrates

The contraction of a species' distribution range, which results from the extirpation of local populations, generally precedes its extinction. Therefore, understanding drivers of range contraction is important for conservation and management. Although there are many processes that can potentially lead to local extirpation and range contraction, three main null models have been proposed: demographic, contagion, and refuge. The first two models postulate that the probability of local extirpation for a given area depends on its relative position within the range; but these models generate distinct spatial predictions because they assume either a ubiquitous (demographic) or a clinal (contagion) distribution of threats. The third model (refuge) postulates that extirpations are determined by the intensity of human impacts, leading to heterogeneous spatial predictions potentially compatible with those made by the other two null models. A few previous studies have explored the generality of some of these null models, but we present here the first comprehensive evaluation of all three models. Using descriptive indices and regression analyses we contrast the predictions made by each of the null models using empirical spatial data describing range contraction in 386 terrestrial vertebrates (mammals, birds, amphibians, and reptiles) distributed across the World. Observed contraction patterns do not consistently conform to the predictions of any of the three models, suggesting that these may not be adequate null models to evaluate range contraction dynamics among terrestrial vertebrates. Instead, our results support alternative null models that account for both relative position and intensity of human impacts. These new models provide a better multifactorial baseline to describe range contraction patterns in vertebrates. This general baseline can be used to explore how additional factors influence contraction, and ultimately extinction for particular areas or species as well as to predict future changes in light of current and new threats.