Constraining null models with environmental gradients: a new method for evaluating the effects of environmental factors and geometric constraints on geographic diversity patterns

The relative effects of climate and geometric constraints on geographic diversity patterns have long been controversial. We developed a new method to assess the role of geometric constraints in shaping altitudinal richness patterns. We showed how plant species richness on four mountains in southwest China are shaped by geometric constraints and environmental gradients together. Contrary to previous studies, our results suggested that: 1) small- and large-ranged species richness were largely controlled by the same environmental gradients, and differed mainly in the effect of geometric constraints. 2) The contribution of geometric constraints (in addition to environmental gradients) to explaining species richness was greater when species richness peaked at low altitudes than at mid-altitudes, suggesting that geometric constraints may be very important when richness peaks are far away from mid-domains. 3) Relating species richness directly to environmental factors (the most widely used method in biodiversity studies) may be misleading when geometric constraints may be affecting the richness pattern, because this method may overestimate the effect of environmental factors by failing to distinguish the confounding effect of geometric constraints. Instead, the effect of environmental factors can be evaluated with an underlying gradient derived from small-ranged species. 4) The geometric constraints effect cannot be fully evaluated by pure geometric constraints models, and is better evaluated with range-based models constrained with environmental gradients. 5) If the generality of our findings is supported for other taxa on other gradients, then many previous studies on the effects of climate and of geometric constraints on geographic diversity patterns may need to be re-visited.