The relative influence of history, climate, topography and vegetation structure on local animal richness varies among taxa and spatial grains.

Understanding the spatial scales at which environmental factors drive species richness patterns is a major challenge in ecology. Due to the trade-off between spatial grain and extent, studies tend to focus on a single spatial scale, and the effects of multiple environmental variables operating across spatial scales on the pattern of local species richness have rarely been investigated. Here, we related variation in local species richness of ground beetles, landbirds and small mammals to variation in vegetation structure and topography, regional climate, biome diversity and glaciation history for 27 sites across the USA at two different spatial grains. We studied the relative influence of broad-scale (landscape) environmental conditions using variables estimated at the site level (climate, productivity, biome diversity and glacial era ice cover) and fine-scale (local) environmental conditions using variables estimated at the plot level (topography and vegetation structure) to explain local species richness. We also examined whether plot-level factors scale up to drive continental scale richness patterns. We used Bayesian hierarchical models and quantified the amount of variance in observed richness that was explained by environmental factors at different spatial scales. For all three animal groups, our models explained much of the variation in local species richness (85%-89%), but site-level variables explained a greater proportion of richness variance than plot-level variables. Temperature was the most important site-level predictor for explaining variance in landbirds and ground beetles richness. Some aspects of vegetation structure were the main plot-level predictors of landbird richness. Environmental predictors generally had poor explanatory power for small mammal richness, while glacial era ice cover was the most important site-level predictor. Relationships between plot-level factors and richness varied greatly among geographical regions and spatial grains, and most relationships did not hold when predictors were scaled up to the continental scale. Our results suggest that the factors that determine richness may be highly dependent on spatial grain, geography, and animal group. We demonstrate that instead of artificially manipulating the resolution to study multiscale effects, a hierarchical approach that uses fine grain data at broad extents could help solve the issue of scale selection in environment-richness studies.