Spatio-temporal metapopulation trends: The coconut crabs of Zanzibar.

Species experience a variety of environmental and anthropogenic conditions across their ranges leading to spatial variation in population dynamics. Understanding population dynamics under different conditions is important but it is challenging to allocate limited effort to spatial and temporal subpopulation monitoring. Using GLMMs, we analyze survey data of a metapopulation of coconut crabs spanning 7 years and 15 sites in and near the Pemba archipelago, Zanzibar, to estimate trends in population size (based on catch per unit effort), weight and sex ratio at the meta- and subpopulation level and investigate anthropogenic drivers of these trends. We found that the overall metapopulation has remained stable in terms of size and composition over the survey period, but observed diverging trends in population size and sex ratio at some subpopulations. Formal protection of sites was associated with positive population trends. Of nine sites for which we could estimate site-specific trends, three showed increasing and two decreasing trends, whereas four sites had stable subpopulations. Although anthropogenic factors affected the average weight, and the incidence of small and large individuals, we found no temporal trends in any weight-related measures. Furthermore, there were no apparent patterns between weight-related measures and subpopulation trends. The metapopulation was biased toward males, and exploitation appeared to be associated with declining trends in the proportion of females, likely an artifact of a strong decline in the proportion of females in one of only two exploited sites in the dataset. Educational campaigns implemented in 2020 at six sites were not related to higher population sizes in later surveys. The variable trends in subpopulation sizes and composition highlight the need for spatially replicated monitoring in metapopulations. The analyses further provide a detailed baseline for future subpopulation studies of this vulnerable species in one of its last remaining metapopulations in the Western Indian Ocean.