Disentangling scale dependencies in species environmental niches and distributions

Understanding species’ responses to environmental conditions, and how these ­species–environment associations shape spatial distributions, are longstanding goals in ecology and biogeography. However, an essential component of species–environment relationships – the spatial unit, or grain, at which they operate – remains unresolved. We identify three components of scale‐dependence in analyses of species–environment associations: 1) response grain, the grain at which species respond most strongly to their environment; 2) environment spatial structure, the pattern of spatial autocorrelation intrinsic to an environmental factor; and 3) analysis grain, the grain at which analyses are conducted and ecological inferences are made. We introduce a novel conceptual framework that defines these scale components in the context of analyzing species–environment relationships, and provide theoretical examples of their interactions for species with various ecological attributes. We then use a virtual species approach to investigate the impacts of each component on common methods of measuring and predicting species–environment relationships. We find that environment spatial structure has a substantial impact on the ability of even simple, univariate species distribution models (SDMs) to recover known species–­environment associations at coarse analysis grains. For simulated environments with ‘fine’ and ‘intermediate’ spatial structure, model explanatory power, and the frequency with which simple SDMs correctly estimated a virtual species’ response to the simulated environment, dramatically declined as analysis grain increased. Informed by these results, we use a scaling analysis to identify maximum analysis grains for individual environmental factors, and a scale optimization procedure to determine the grain of maximum predictive accuracy. Implementing these analysis grain thresholds and model performance standards in an example east African study system yields more accurate distribution predictions, compared to SDMs independently constructed at arbitrary analysis grains. Finally, we integrate our conceptual framework with virtual and empirical results to provide practical recommendations for researchers asking common questions about species–environment relationships.