Abstract
Mark-recapture estimators are commonly used for population size estimation, and typically yield unbiased estimates for most solitary species with low to moderate home range sizes. However, these methods assume independence of captures among individuals, an assumption that is clearly violated in social species that show fission-fusion dynamics, such as the Asian elephant. In the specific case of Asian elephants, doubts have been raised about the accuracy of population size estimates. More importantly, the potential problem for the use of mark-recapture methods posed by social organization in general has not been systematically addressed. We developed an individual-based simulation framework to systematically examine the potential effects of type of social organization, as well as other factors such as trap density and arrangement, spatial scale of sampling, and population density, on bias in population sizes estimated by POPAN, Robust Design, and Robust Design with detection heterogeneity. In the present study, we ran simulations with biological, demographic and ecological parameters relevant to Asian elephant populations, but the simulation framework is easily extended to address questions relevant to other social species. We collected capture history data from the simulations, and used those data to test for bias in population size estimation. Social organization significantly affected bias in most analyses, but the effect sizes were variable, depending on other factors. Social organization tended to introduce large bias when trap arrangement was uniform and sampling effort was low. POPAN clearly outperformed the two Robust Design models we tested, yielding close to zero bias if traps were arranged at random in the study area, and when population density and trap density were not too low. Social organization did not have a major effect on bias for these parameter combinations at which POPAN gave more or less unbiased population size estimates. Therefore, the effect of social organization on bias in population estimation could be removed by using POPAN with specific parameter combinations, to obtain population size estimates in a social species.
Highlights
Population size estimation is a vital aspect of studying the ecology of animal populations in the wild
While we carried out this study with the specific aim of assessing commonly used mark-recapture methods in the context of population size estimation for Asian elephant populations in the wild, we expect that the simulation framework we have developed will be applicable to many other species of medium to large sized mammals with relatively minor modifications
The association indices (AIs) distribution data indicate that the different patterns of Association Probabilities (APs) we used in our simulations did, result in AI distributions reflecting the three social organizations intended
Summary
Population size estimation is a vital aspect of studying the ecology of animal populations in the wild. The problem is that markrecapture methods typically assume the detection probabilities of individuals to be independent [8,9,10], whereas this assumption is likely to be violated in social species that show coordinated movement of socially interacting individuals The effects of such non-independence of capture probabilities can be non-trivial: Boulanger et al [11] found that non-independence between pairs of individuals (mother-offspring pairs in their case) caused population size estimates to be biased, despite incorporating heterogeneity in capture probabilities between those individuals in their simulation. The capture of entire groups rather than individuals is considered, and the estimate of total groups can be multiplied by the mean group size to get an estimate of total population size [8] In cases where this approach cannot be used, corrections need to be made for the variance of mark-recapture parameters. More sophisticated numerical methods are called for in such cases [13,15]
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