The shift between the Red Queen and the Red King effects in mutualisms.

Interspecific mutualisms consist of partners trading services that yield common benefits to both species. Until now, understanding how the payoffs from mutualistic cooperation are allocated among the participants has been problematic. Two hypotheses have been proposed to resolve this problem. The Red Queen effect argues that faster-evolving species are favoured in co-evolutionary processes because they are able to obtain a larger share of benefits. Conversely, the Red King effect argues that the slower-evolving species gains a larger share of benefits. The model we propose shows that the allocations for a common benefit vary when the effect of a reward mechanism is included in the model. The outcome is a shift from the Red Queen effect to the Red King effect and vice versa. In addition, our model shows that either an asymmetry in payoff or an asymmetry in the number of cooperative partners causes a shift between the Red Queen effect and the Red King effect. Even in situations where the evolutionary rates are equal between the two species, asymmetries in rewards and in participant number lead to an uneven allocation of benefits among the partners.