Ant diversity in Neotropical savannas: Hierarchical processes acting at multiple spatial scales.

Understanding what creates and maintains macroscale biodiversity gradients is a central focus of ecological and evolutionary research. Spatial patterns in diversity are driven by a hierarchy of factors operating at multiple scales. Historical and climatic factors drive large-scale patterns of diversity by affecting the size of regional species pools, while habitat heterogeneity or microhabitat characteristics further influence species coexistence at small scales. We tested the degree to which the species-energy, historical factors, habitat heterogeneity and local environment hypotheses explain observed patterns of ant diversity across hierarchical spatial scales. We sampled ground-dwelling ants at 29 sites within a Neotropical savanna region, the Brazilian Cerrado. We measured species density - an abundance-dependent diversity metric - and rarefied species richness - an abundance-independent metric - at spatial scales with varying grain sizes. For each hypothesis, two correlates were used to predict ant diversity patterns: (a) species-energy: rainfall and productivity; (b) historical factors: historical variation in rainfall and refugial areas; (c) habitat heterogeneity: heterogeneity in greenness and diversity of land cover; and (d) local factors: contents of sand and coarse fragments in the soil. Ant diversity patterns correlated to net primary productivity and to the proportion of coarse fragments in the soil, corroborating the species-energy and local environment hypotheses, respectively. Soil negatively influenced species density, but not rarefied species richness, which was positively influenced by productivity. We found scale dependencies in the effects of soil/productivity on species density; productivity best predicted species density patterns at large scales, since sampling completeness offset the abundance-driven effects of soil. Considering abundance differences may help to discern the mechanisms underlying the relationship between macroscale diversity patterns and its ecological drivers. Plant productivity affected ant diversity independently of abundance, possibly by limiting the size of regional species pools. On the other hand, soil properties had an abundance-dependent effect on ant diversity, indicating a sampling mechanism. Our findings are consistent with predictions of the hierarchical theory of diversity. Large-scale patterns of productivity limit regional diversity, an effect that cascades down to finer spatial scales, where soil properties influence the number of coexisting species.