Small-mammal density estimation: A field comparison of grid-based vs. web-based density estimators

Statistical models for estimating absolute densities of field populations of animals have been widely used over the last century in both scientific studies and wildlife management programs. To date, two general classes of density estimation models have been developed: models that use data sets from capture-recapture or removal sampling techniques (often derived from trapping grids) from which separate estimates of population size ((N) over cap) and effective sampling area ((A) over cap) are used to calculate density ((D) over cap = (N) over cap/(A) over cap); and models applicable to sampling regimes using distance-sampling theory (typically transect lines or trapping webs) to estimate detection functions and densities directly from the distance data. However, few studies have evaluated these respective models for accuracy, precision. and bias on known field populations. and no studies have been conducted that compare the two approaches under controlled field conditions. In this study, we evaluated both classes of density estimators on known densities of enclosed rodent populations. Test data sets (n = 11) were developed using nine rodent species from capture-recapture live-trapping on both trapping grids and trapping webs in four replicate 4.2-ha enclosures on the Sevilleta National Wildlife Refuge in central New Mexico, USA. Additional "saturation" trapping efforts resulted in an enumeration of the rodent populations in each enclosure, allowing the computation of true densities. Density estimates ((D) over cap) were calculated using program CAPTURE for the grid data sets and program DISTANCE for the web data sets, and these results were compared to the known true densities (D) to evaluate each model's relative mean square error, accuracy, precision, and bias. In addition, we evaluated a variety of approaches to each data set's analysis by having a group of independent expert analysts calculate their best density estimates without a priori knowledge of the true densities this "blind" test allowed us to evaluate the influence of expertise and experience in calculating density estimates in comparison to simply using default values in programs CAPTURE and DISTANCE. While the rodent sample sizes were considerably smaller than the recommended minimum for good model results, we found that several models performed well empirically. including the web-based uniform and half-normal models in program DISTANCE, and the grid-based models M-b and M-bh in program CAPTURE (with A adjusted by species-specific full mean maximum distance moved (MMDM) values). These models produced accurate (D) over cap values (with 95% confidence intervals that included the true D values) and exhibited acceptable bias but poor precision. However. in linear regression analyses comparing each model's (D) over cap values to the true D values over the range of observed test densities, only the web-based uniform model exhibited a regression slope near 1.0; all other models showed substantial slope deviations. indicating biased estimates at higher or lower density values. In addition, the grid-based (D) over cap analyses using full MMDM values for (W) over cap area adjustments required a number of theoretical assumptions of uncertain validity, and we therefore viewed their empirical successes with caution. Finally, density estimates from the independent analysts were highly variable, but estimates from web-based approaches had smaller mean square errors and better achieved confidence-interval coverage of D than did grid-based approaches.

Our results support the contention that web-based approaches for density estimation of small-mammal populations are both theoretically and empirically superior to grid-based approaches, even when sample size is far less than often recommended. In view of the increasing need for standardized environmental measures for comparisons among ecosystems and through time, analytical models based on distance sampling appear to offer accurate density estimation approaches for research studies involving small-mammal abundances.