Drivers of distributions and niches of North American cold-adapted amphibians: evaluating both climate and land use.

Species distribution estimates are often used to understand the niche of a species; however, these are often based solely on climatic predictors. When the influences of biotic factors are ignored, erroneous inferences about range and niche may be made. We aimed to integrate climate data with a unique set of available land cover and land use data for the six cold-adapted amphibians of North America (Ambystoma macrodactylum, Anaxyrus hemiophrys, Anaxyrus boreas, Pseudacris maculata, Rana sylvatica, Rana luteiventris) to determine the relative importance of climate and non-climate drivers through the use of ecological niche models for present-day range estimates. We compared climate-only, land use-only, and combination models of climate and land use, derived from two different model selection techniques, to determine which was most likely to drive current distributions of cold-adapted amphibian species. Land use layers included land cover type, human population, vegetation type, ecoregion, and the overall human footprint. The most supported models included both climate and land use, with climate and human footprint variables having the highest permutation importance and percent contribution. Models that incorporated climate and land use data performed best as measured with AIC and AUC, although qualitatively most underestimated the northern range edge, implying potential sampling bias or locations of reduced habitat quality for these species in the northern area of the ranges. There were small differences in overall combination models dependent on the method of model selection. The overall effect sizes of landscape factors within the combination models were small except for one landscape feature: human footprint, which incorporated multiple aspects of anthropogenic change on the landscape, including human population density, travel access, and agricultural impact. This aspect of the landscape was just as important as climate, and counter to what we expected, the association was mostly positive, with a negative response only occurring at very high levels. This highlights the importance of moving beyond climate only species range estimates as land cover, specifically human impact, may be driving the patterns of species' ranges.