Where do they go? The effects of topography and habitat diversity on reducing climatic debt in birds.

The spatial tracking of climatic shifts is frequently reported as a biodiversity response to climatic change. However, species' range shifts are often idiosyncratic and inconsistent with climatic shift predictions. At the community scale, this discrepancy can be measured by comparing the spatial shift in the relative composition of cold- vs. warm-adapted species in a local assemblage [the community temperature index (CTI)] with the spatial shift in temperature isotherms. While the local distribution of climate change velocity is a promising approach to downscaling climate change pressure and responses, CTI velocity has only been investigated on a continental or national scale. In this study, we coupled French Breeding Bird Survey data, collected from 2133 sites monitored between 2001 and 2012, with climatic data in order to estimate the local magnitude and direction of breeding season temperature shift, CTI shift, and their spatiotemporal divergence - the local climatic debt. We also tested whether landscape characteristics that are known to affect climate velocity and spatial tracking of climate change mediated the climatic debt on the local scale. We found a clear spatial structure, together with heterogeneity in both temperature and CTI spatial shifts. Local climatic debt decreased as the elevation, habitat diversity, and the naturalness of the landscape increased. These results suggest the complementary effects of the local topographic patterns sheltering more diverse microclimates and the increasing permeability of natural and diversified landscape. Our findings suggest that a more nuanced evaluation of spatial variability in climatic and biotic shifts is necessary in order to properly describe biodiversity responses to climate change rather than the oversimplified descriptions of uniform poleward shifts.