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Sex-specific population dynamics and demography of capercaillie (Tetrao urogallus L.) in a patchy environment

Modeling the population dynamics of patchily distributed species is a challenge, particularly when inference must be based on incomplete and small data sets such as those from most species of conservation concern. Open population spatial capture-recapture (SCR) models are ideally suited to quantify population trends, but have seen only limited use since their introduction. To investigate population trend and sex-specific population dynamics, we applied an open SCR model to a capercaillie (Tetrao urogallus) population in Switzerland living in eight distinct forest patches totalling 22 km2 within a region of 908 km2. The population was surveyed using genetic sampling of scat in 2009, 2012 and 2015. We fit an open SCR model with sex-specific detection and population dynamics parameters while accounting for the patchy distribution of habitat and the uncertainty introduced by observing the population in three years only. Between 2009 and 2015, a total of 143 males, 112 females and 4 individuals of uncertain sex were detected. The annual per capita recruitment rate was estimated at 0.115 (SE 0.0144) for males and 0.127 (0.0168) for females. The estimated annual survival probability for males was 0.758 (0.0241) and 0.707 (0.0356) for females. The population trajectory implied by these survival and recruitment estimates was a decline of 2% per year; however, the sex specificity of the model revealed a decline in the male population only, with no evidence of decline in the female population. Further, the population decline observed in males was explained by the demography of just two of the eight patches. Using a customized open population SCR model, we determined that the endangered capercaillie in our Swiss study area had a stable female population and a declining male population, with the male decline due to population dynamics in a subset of the study area. Our study highlights the flexibility of open population SCR models for assessing population trajectories through time and across space and emphasizes the desirability of estimating sex-stratified population trends especially in species of conservation concern.