Discriminative host sanction together with relatedness promote the cooperation in fig/fig wasp mutualism

Sanctioning or punishing is regarded as one of the most important dynamics in the evolution of cooperation. However, it has not been empirically examined yet whether or not such enforcement selection by sanctioning or punishing and classical theories like kin or reciprocity selection are separate mechanisms contributing to the evolution of cooperation. In addition, it remains largely unclear what factors determine the intensity or effectiveness of sanction. Here, we show that in the obligate, interspecific cooperation between figs and fig wasps, the hosted figs can discriminatively sanction cheating individuals by decreasing the offspring development ratio. Concurrently, the figs can reward the cooperative pollinators with a higher offspring development ratio. This sanction intensity and effectiveness largely depend on how closely the host and symbiont are related either in terms of reciprocity exchange or genetic similarity as measured by the reciprocal of the foundress number. Our results imply that in asymmetric systems, symbionts might be forced to evolve to be cooperative or even altruistic through discriminative sanction against the noncooperative symbiont and reward to the cooperative symbiont by the host (i.e. through a game of 'carrot and stick').