Steady-State and Dynamical Behavior of a PDE Model of Multilevel Selection with Pairwise Group-Level Competition.

Evolutionary competition often occurs simultaneously at multiple levels of organization, in which traits or behaviors that are costly for an individual can provide collective benefits to groups to which the individual belongs. Building off of recent work that has used ideas from game theory to study evolutionary competition within and among groups, we study a PDE model for multilevel selection that considers group-level evolutionary dynamics through a pairwise conflict depending on the strategic composition of the competing groups. This model allows for incorporation of group-level frequency dependence, facilitating the exploration for how the form of probabilities for victory in a group-level conflict can impact the long-time support for cooperation via multilevel selection. We characterize well-posedness properties for measure-valued solutions of our PDE model and apply these properties to show that the population will converge to a delta-function at the all-defector equilibrium when between-group selection is sufficiently weak. We further provide necessary conditions for the existence of bounded steady state densities for the multilevel dynamics of Prisoners' Dilemma and Hawk-Dove scenarios, using a mix of analytical and numerical techniques to characterize the relative strength of between-group selection required to ensure the long-time survival of cooperation via multilevel selection. We also see that the average payoff at steady state appears to be limited by the average payoff of the all-cooperator group, even for games in which groups achieve maximal average payoff at intermediate levels of cooperation, generalizing behavior that has previously been observed in PDE models of multilevel selection with frequency-independent group-level competition.