Does your species have memory? Analyzing capture–recapture data with memory models

We examine memory models for multisite capture-recapture data. This is an important topic, as animals may exhibit behavior that is more complex than simple first-order Markov movement between sites, when it is necessary to devise and fit appropriate models to data. We consider the Arnason-Schwarz model for multisite capture-recapture data, which incorporates just first-order Markov movement, and also two alternative models that allow for memory, the Brownie model and the Pradel model. We use simulation to compare two alternative tests which may be undertaken to determine whether models for multisite capture-recapture data need to incorporate memory. Increasing the complexity of models runs the risk of introducing parameters that cannot be estimated, irrespective of how much data are collected, a feature which is known as parameter redundancy. Rouan et al. (JABES, 2009, pp 338-355) suggest a constraint that may be applied to overcome parameter redundancy when it is present in multisite memory models. For this case, we apply symbolic methods to derive a simpler constraint, which allows more parameters to be estimated, and give general results not limited to a particular configuration. We also consider the effect sparse data can have on parameter redundancy and recommend minimum sample sizes. Memory models for multisite capture-recapture data can be highly complex and difficult to fit to data. We emphasize the importance of a structured approach to modeling such data, by considering a priori which parameters can be estimated, which constraints are needed in order for estimation to take place, and how much data need to be collected. We also give guidance on the amount of data needed to use two alternative families of tests for whether models for multisite capture-recapture data need to incorporate memory.