Here, we consider a game-theoretic model of multilevel selection in which individuals compete based on their payoff and groups also compete based on the average payoff of group members. Our focus is on multilevel social dilemmas: games in which individuals are best off cheating, while groups of individuals do best when composed of many cooperators. We analyze the dynamics of the two-level replicator dynamics, a nonlocal hyperbolic PDE describing deterministic birth-death dynamics for both individuals and groups. While past work on such multilevel dynamics has restricted attention to scenarios with exactly solvable within-group dynamics, we use comparison principles and an invariant property of the tail of the population distribution to extend our analysis to all possible two-player, two-strategy social dilemmas. In the Stag-Hunt and similar games with coordination thresholds, we show that any amount of between-group competition allows for fixation of cooperation in the population. For the prisoners' dilemma and Hawk-Dove game, we characterize the threshold level of between-group selection dividing a regime in which the population converges to a delta function at the equilibrium of the within-group dynamics from a regime in which between-group competition facilitates the existence of steady-state densities supporting greater levels of cooperation. In particular, we see that the threshold selection strength and average payoff at steady state depend on a tug-of-war between the individual-level incentive to be a defector in a many-cooperator group and the group-level incentive to have many cooperators over many defectors. We also find that lower-level selection casts a long shadow: If groups are best off with a mix of cooperators and defectors, then there will always be fewer cooperators than optimal at steady state, even in the limit of infinitely strong competition between groups.
Keywords: Evolutionary game theory; Multilevel selection; Replicator equations.