Ideal free distributions, evolutionary games, and population dynamics in multiple-species environments

In this article, we develop population game theory, a theory that combines the dynamics of animal behavior with population dynamics. In particular, we study interaction and distribution of two species in a two-patch environment assuming that individuals behave adaptively (i.e., they maximize Darwinian fitness). Either the two species are competing for resources or they are in a predatorprey relationship. Using some recent advances in evolutionary game theory, we extend the classical ideal free distribution (IFD) concept for single species to two interacting species. We study population dynamical consequences of two-species IFD by comparing two systems: one where individuals cannot migrate between habitats and one where migration is possible. For single species, predator-prey interactions, and competing species, we show that these two types of behavior lead to the same population equilibria and corresponding species spatial distributions, provided interspecific competition is patch independent. However, if differences between patches are such that competition is patch dependent, then our predictions strongly depend on whether animals can migrate or not. In particular, we show that when species are settled at their equilibrium population densities in both habitats in the environment where migration between habitats is blocked, then the corresponding species spatial distribution need not be an IFD. Thus, when species are given the opportunity to migrate, they will redistribute to reach an IFD (e.g., under which the two species can completely segregate), and this redistribution will also influence species population equilibrial densities. Alternatively, we also show that when two species are distributed according to the IFD, the corresponding population equilibrium can be unstable.