Abstract
Aims Studies of species distribution patterns traditionally have been conducted at a single scale, often overlooking species-environ- ment relationships operating at finer or coarser scales. Testing diversity-related hypotheses at multiple scales requires a robust sampling design that is nested across scales. Our chief motivation in this study was to quantify the contributions of different predictors of herbaceous species richness at a range of local scales. Methods Here, we develop a hierarchically nested sampling design that is balanced across scales, in order to study the role of several envi- ronmental factors in determining herbaceous species distribution at various scales simultaneously. We focus on the impact of woody vegetation, a relatively unexplored factor, as well as that of soil and topography. Light detection and ranging (LiDAR) imaging enabled precise characterization of the 3D structure of the woody veg- etation, while acoustic spectrophotometry allowed a particularly high-resolution mapping of soil CaCO3 and organic matter contents. Important Findings We found that woody vegetation was the dominant explanatory variable at all three scales (10, 100 and 1 000 m 2 ), accounting for more than 60% of the total explained variance. In addition, we found that the species richness-environment relationship was scale dependent. Many studies that explicitly address the issue of scale do so by comparing local and regional scales. Our results show that efforts to conserve plant communities should take into account scale dependence when analyzing species richness-envi- ronment relationships, even at much finer resolutions than local vs. regional. In addition, conserving heterogeneity in woody veg- etation structure at multiple scales is a key to conserving diverse herbaceous communities.
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