Anthropogenic and environmental drivers of modern range loss in large mammals.

The extinction of a species is inevitably preceded by the extirpation of a series of local populations. Ecological theory predicts that vulnerability to extirpation varies between populations and is ultimately linked to environmental heterogeneity. If populations of a species are present in multiple regions separated by abrupt changes in environmental conditions (e.g., biomes), spatial variation in vulnerability to extirpation may be closely linked to the distribution of these regions. In the absence of abrupt shifts in environmental conditions, populations at the edge of a species' range should have low growth rates and be more vulnerable to extirpation, whereas populations located in the core of the species' range should be exposed to more favorable environmental conditions, have higher growth rates, and be less vulnerable. Here, we ask whether the distribution of biomes or range position better reflects spatial variation in vulnerability for 43 mammal species distributed through four continents. We control for the distribution of human threats and quantify the importance of protected areas in population persistence. We conclude that the distribution of biomes is a better predictor of vulnerability than position in the geographic range. We also find that core populations are less vulnerable than edge populations (after controlling for threats levels and protected areas). Protected areas are important for the persistence of most species we studied. By providing a measure of vulnerability linked directly to the distribution of threats, our results offer insights for scaling up from species vulnerability to extinction risk.