Estimating true instead of apparent survival using spatial Cormack-Jolly-Seber models

Survival is often estimated from capture-recapture data using Cormack-Jolly-Seber (CJS) models, where mortality and emigration cannot be distinguished, and the estimated apparent survival probability is the product of the probabilities of true survival and of study area fidelity. Consequently, apparent survival is lower than true survival unless study area fidelity equals one. Underestimation of true survival from capture-recapture data is a main limitation of the method. We develop a spatial version of the CJS model that allows estimation of true survival. Besides the information about whether a specific individual was encountered at a given occasion, it is often recorded where the encounter occurred. Thus, information is available about the fraction of dispersal that occurs within the study area, and we use it to model dispersal and estimate true survival. Our model is formulated hierarchically and consists of survival, dispersal and observation submodels, assuming that encounters are possible anywhere within a study area. In a simulation study, our new spatial CJS model produced accurate estimates of true survival and dispersal behaviour for various sizes and shapes of the study area, even if emigration is substantial. However, when the information about dispersal is scarce due to low survival, low recapture probabilities and high emigration, the estimators are positively biased. Moreover, survival estimates are sensitive to the assumed dispersal kernel. We applied the spatial CJS model to a data set of adult red-backed shrikes (Lanius collurio). Apparent survival of males (c.05) estimated with the CJS model was larger than in females (c.04), but the application of the spatial CJS model revealed that both sexes had similar survival probabilities (c.06). The mean breeding dispersal distance in females was c.700m, while males dispersed only c.250m between years. Spatial CJS models enable study of dispersal and survival independent of study design constraints such as imperfect detection and size of the study area provided that some of the dispersing individuals remain in the study area. We discuss possible extensions of our model: alternative dispersal models and the inclusion of covariates and of a habitat suitability map.