Multistate survival models and their extensions in Program MARK

Program MARK provides .100 models for the estimation of population parameters from mark?encounter data. The multistate model of Brownie et al. (1993) and Hestbeck et al. (1991) allows animals to move between states with a probability of transition. The simplest multistate model is an extension of the Cormack?Jolly?Seber (CJS) live recapture model. arameters estimated are state-specific survival rates and encounter probabilities and transition probabilities between states. The multistate model provides a valuable framework to evaluate important ecological questions. For example, estimation of state-specific survival and transition probabilities between the biological states of breeders and nonbreeders allows estimation of the cost of reproduction. Transitions between physical states, such as spatial areas, provide estimates needed for meta-population models. The basic multistate model uses only live recaptures, but 3 extensions are included in MARK. A multistate model with live and dead encounters is available, although the dead encounters are not state specific. Robust-design multistate models are also included in MARK, with both open and closed robust designs. These models assume that animals move between states only between primary sessions of the robust design. For the closed robust design, we can specify 12 different data types for the modeling of encounter probabilities during the primary session, including 6 versions of the closed model likelihood incorporating population size (N) directly in the likelihood, and 6 versions of the Huggins model in which N is estimated as a derived parameter outside the likelihood. One assumption that is generally necessary to estimate state-specific survival rates in the multistate model is that transitions take place immediately before encounter occasions. Otherwise, survival rates over the interval between encounter occasions are a mix of survival rates over multiple states. Advantages of using MARK to estimate the parameters of the various multistate models include flexibility of model specification to include group, time, and individual covariates, estimation of variance components, model averaging of parameter estimates, and Bayesian parameter estimation using Markov chain Monte Carlo procedures on the logit scale.