Estimating animal abundance at multiple scales by spatially explicit capture-recapture.

Information about how animal abundance varies across landscapes is needed to inform management action but is costly and time-consuming to obtain; surveys of a single population distributed over a large area can take years to complete. Surveys employing small, spatially replicated sampling units improve efficiency, but statistical estimators rely on assumptions that constrain survey design or become less reasonable as larger areas are sampled. Efficient methods that avoid assumptions about similarity of detectability or density among replicates are therefore appealing. Using simulations and data from >3500 black bears sampled on 73 independent study areas in Ontario, Canada, we (1) quantified bias induced by unmodeled spatial heterogeneity in detectability and density; (2) evaluated novel, design-based estimators of average density across replicate study areas; and (3) evaluated two estimators of the variance of average density across study areas: an analytic estimator that assumed an underlying homogeneous spatial Poisson point process for the distribution of animals' activity centers, and an empirical estimator of variance across study areas. In simulations where detectability varied in space, assuming spatially constant detectability yielded density estimates that were negatively biased by 20% to 30%; estimating local detectability and density from local data and treating study areas as independent, equal replicates when estimating average density across study areas using the design-based estimator yielded unbiased estimates at local and landscape scales. Similarly, detectability of black bears varied among study areas and estimates of bear density at landscape scales were higher when no information was shared across study areas when estimating detectability. This approach also maximized precision (relative SEs of estimates of average black bear density ranged from 7% to 18%) and computational efficiency. In simulations, the analytic variance estimator was robust to threefold variation in local densities but the empirical estimator performed poorly. Conducting multiple, similar SECR surveys and treating them as independent replicates during analyses allowed us to efficiently estimate density at multiple scales and extents while avoiding biases caused by pooling spatially heterogeneous data. This approach enables researchers to address a wide range of ecological or management-related questions and is applicable with most types of SECR data.