Elements of regional beetle faunas: faunal variation and compositional breakpoints along climate, land cover and geographical gradients.

Regional faunas are structured by historical, spatial and environmental factors. We studied large-scale variation in four ecologically different beetle groups (Coleoptera: Dytiscidae, Carabidae, Hydrophiloidea, Cerambycidae) along climate, land cover and geographical gradients, examined faunal breakpoints in relation to environmental variables, and investigated the best fit pattern of assemblage variation (i.e. randomness, checkerboards, nestedness, evenly spaced, Gleasonian, Clementsian). We applied statistical methods typically used in the analysis of local ecological communities to provide novel insights into faunal compositional patterns at large spatial grain and geographical extent. We found that spatially structured variation in climate and land cover accounted for most variation in each beetle group in partial redundancy analyses, whereas the individual effect of each explanatory variable group was generally much less important in accounting for variation in provincial species composition. We also found that climate variables were most strongly associated with faunal breakpoints, with temperature-related variables alone accounting for about 20% of variation at the first node of multivariate regression tree for each beetle group. The existence of faunal breakpoints was also shown by the 'elements of faunal structure' analyses, which suggested Clementsian gradients across the provinces, that is, that there were two or more clear groups of species responding similarly to the underlying ecological gradients. The four beetle groups showed highly similar biogeographical patterns across our study area. The fact that temperature was related to faunal breakpoints in the species composition of each beetle group suggests that climate sets a strong filter to the distributions of species at this combination of spatial grain and spatial extent. This finding held true despite the ecological differences among the four beetle groups, ranging from fully aquatic to fully terrestrial and from herbivorous to predaceous species. The existence of Clementsian gradients may be a common phenomenon at large scales, and it is likely to be caused by crossing multiple species pools determined by climatic and historical factors on the distributions of species.