Counting Sunda clouded leopards with confidence: incorporating individual heterogeneity in density estimates

Even with intensive sampling effort, data often remain sparse when estimating population density of elusive species such as the Sunda clouded leopard Neofelis diardi. An inadequate number of recaptures can make it difficult to account for heterogeneity in detection parameters. We used data from large-scale camera-trapping surveys in three forest reserves in Sabah, Malaysian Borneo, to (1) examine whether a high-density camera-trap network increases the number of recaptures for females, which tend to be more difficult to detect, thus improving the accuracy of density estimates; (2) compare density estimates from models incorporating individual heterogeneity in detection parameters with estimates from the null model to evaluate its potential bias; and (3) investigate how the size of the camera-trap grid affects density and movement estimates. We found that individual heterogeneity could not be incorporated in the single-site data analysis and only conservative null model estimates could be generated. However, aggregating data across study sites enabled us to account for individual heterogeneity and we estimated densities of 1.27-2.82 individuals/100 km(2), 2-3 times higher than estimates from null models. In light of these findings, it is possible that earlier studies underestimated population density. Similar densities found in well-managed forest and recently selectively logged forest suggest that Sunda clouded leopards are relatively resilient to forest disturbances. Our analysis also revealed that camera-trapping grids for Sunda clouded leopard density estimations should cover large areas (c. 250 km(2)), although smaller grids could be appropriate if density or detectability are higher.