Sanctions, partner recognition, and variation in mutualism

Mutualistic interactions can be stabilized against invasion by noncooperative individuals by putting such “cheaters” at a selective disadvantage. Selection against cheaters should eliminate genetic variation in partner quality—yet such variation is often found in natural populations. One explanation for this paradox is that mutualism outcomes are determined not only by responses to partner performance but also by partner signals. Here, we build a model of coevolution in a symbiotic mutualism, in which hosts’ ability to sanction noncooperative symbionts and recognition of symbiont signals are determined by separate loci, as are symbionts’ cooperation and expression of signals. In the model, variation persists without destabilizing the interaction, in part because coevolution of symbiont signals and host recognition is altered by the coevolution of sanctions and cooperation, and vice versa. Individual-based simulations incorporating population structure strongly corroborate these results....