Abstract
Predicting the outcomes of land management on biodiversity is difficult without a mechanistic understanding of how management approaches, ecosystem structure, environmental conditions, and biodiversity interact. Management effects may be direct or indirect, context- or scale-dependent, or obscured by local environmental conditions. Resolving these relationships at the regional scale may be difficult, given heterogeneity in local environmental conditions, yet understanding broad-scale patterns can elucidate context dependencies and improve restoration outcomes. We confronted these challenges within globally rare oak savannas in the midwestern United States, which have been altered by fire exclusion and resulting woody encroachment. By modeling direct and indirect pathways by which management influences diversity, we test a general framework for savanna restoration. Across 100 oak savannas spanning five US states, management by prescribed fire and mechanical thinning of woody vegetation affected groundlayer plant species richness through changes to ecosystem structure (canopy openness and litter depth), and these effects were both context- and scale-dependent. Frequent prescribed fires and canopy thinning promoted greater canopy openness, which in turn increased richness at small (1 m(2)), but not larger (1000 m(2)) scales. Frequent fire additionally increased richness at small and larger scales through effects independent of ecosystem structure. While management effects were large relative to the influence of local edaphic conditions, soil productivity had two largely offsetting effects on small-scale richness, increasing richness directly but decreasing richness indirectly by promoting closed canopy structure. These results suggest using a combination of fire and canopy thinning to reverse the effects of decades of fire exclusion. However, management effects were also context-dependent, emphasizing that management outcomes vary regionally. Here, 1-m(2) plant species richness increased with both fire frequency and canopy thinning under low, but not high, productivity soil conditions. By demonstrating how specific management practices influence savanna structure and biodiversity by manipulating ecological processes across broad geographic and edaphic gradients, our findings provide a framework for understanding management outcomes at short and medium intervals (e.g., within and between decades, respectively), in the form of a model that can be refined by testing additional hypotheses to better predict savanna restoration outcomes.