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.

Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home-range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied block cross-validation to quantify bias in empirical home-range estimates. Area requirements of mammals <10 kg were underestimated by a mean approximately15%, and species weighing approximately100 kg were underestimated by approximately50% on average. Thus, we found area estimation was subject to autocorrelation-induced bias that was worse for large species. Combined with the fact that extinction risk increases as body mass increases, the allometric scaling of bias we observed suggests the most threatened species are also likely to be those with the least accurate home-range estimates. As a correction, we tested whether data thinning or autocorrelation-informed home-range estimation minimized the scaling effect of autocorrelation on area estimates. Data thinning required an approximately93% data loss to achieve statistical independence with 95% confidence and was, therefore, not a viable solution. In contrast, autocorrelation-informed home-range estimation resulted in consistently accurate estimates irrespective of mass. When relating body mass to home range size, we detected that correcting for autocorrelation resulted in a scaling exponent significantly >1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum.

One of the greatest challenges in restoring species to the wild is insufficient knowledge about their habitat requirements and movement ecology. This is especially true for wide-ranging species such as the scimitar-horned oryx (Oryx dammah). Once widespread across Sahelo-Saharan grasslands, oryx were declared Extinct in the Wild in 1999. Here, we integrate GPS/satellite tracking, remote sensing, and movement analyses to assess how reintroduced oryx respond to wild conditions. We monitored two groups of oryx, reared under different captive management regimes and released in different seasons, for 12 months after release. Our study provides the first movement trajectories and home range estimates for this species. We expected oryx movements after release to represent trade-offs between risky, energetically expensive exploration and resource exploitation. Oryx raised under semi-free ranging conditions and released during the wet season ("ranging") exhibited this pattern of exploration followed by home range establishment. In contrast, oryx raised in small pens and released during the dry season ("penned") explored far less novel terrain. Ranging oryx exhibited seasonal shifts in activity and movement timing, while penned oryx simply reduced overall movement and continuously accessed supplemental food and water. Sahelian ecosystems exhibit strong seasonal cycles and extensive spatial variation. In this highly variable environment, reintroduced oryx will need to disperse from the release site to acquire adequate forage throughout the year. Thus, we experimentally varied acclimation period, and expected dispersal to decrease with acclimation period length. Post-release dispersal ranged from 2 to 90 km: ranging oryx acclimated for ca. 6 months moved 40-60 km from the release site, while penned oryx acclimated for ca. 1 month remained within 5-25 km. Our results demonstrate that captive management and environmental conditions at release strongly influence the extent to which reintroduced oryx disperse and adapt to wild conditions. We also show that-in contrast to previous studies-longer acclimation periods do not necessarily lead to site fidelity. Finally, our findings demonstrate the importance of tracking a large proportion of reintroduced individuals to (1) accurately record post-release behaviors and vital rates, and (2) adaptively evaluate pre-and post-release management actions to improve conservation outcomes.

The characterization of species’ environmental niches and spatial distribution predictions based on them are now central to much of ecology and conservation, but implicitly requires decisions about the appropriate spatial scale (i.e., grain) of analysis. Ecological theory and empirical evidence suggest that range‐resident species respond to their environment at two characteristic, hierarchical spatial grains: (1) response grain, the (relatively fine) grain at which an individual uses environmental resources, and (2) occupancy grain, the (relatively coarse) grain equivalent to a typical home range. We use a multi‐grain (MG) occupancy model, aided by fine‐grain remotely sensed imagery, to simultaneously estimate species–environment associations at both grains, conduct grain optimization to measure response grain, and apply this analysis framework to an example species: a medium‐sized bird (Tockus deckeni) in a heterogeneous East African landscape. Based on home range analysis of movement data, we calculate an occupancy grain of 1 km for T. deckeni. Using a grain optimization procedure across 32 grains from 10 to 500 m, we identify 60 m as the most strongly supported response grain for a suite of environmental variables, slightly coarser than opportunistic behavioral observations would have suggested. Validation confirms that the accuracy of the optimized MG occupancy model substantially exceeds that of equivalent single‐grain (SG) occupancy models. We further use a simulation approach to assess the potential impacts of accounting for the multi‐scale structure of species’ environmental requirements on estimates of population size. We find that the more strongly supported MG approach consistently predicts a minimum population size in the study landscape that is much lower than that provided by the SG model. This suggests that SG approaches commonly used in conservation applications could lead to overly optimistic abundance and population estimates, and that the MG approach may be more appropriate for supporting species conservation goals. More generally, we conclude that multi‐grain approaches of the sort presented, and increasingly enabled by growing high‐resolution remotely sensed data, hold great promise for offering a more mechanistic framework for assessing the appropriate grain(s) for population monitoring and management and enable more reliable estimates of abundances and species’ distributions.