A missing model in reproductive skew theory: The bordered tug-of-war

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Abstract
Models of reproductive skew can be classified into two groups: transactional models, in which group members yield shares of reproduction to each other in return for cooperation, and tug-of-war models, in which group members invest group resources in a tug-of-war over their respective reproductive shares. We synthesize these two models to yield a "bordered tug-of-war" model in which the internal tug-of-war is limited ("bordered") by the requirement that group members must achieve a certain amount of reproduction lest they pursue a noncooperative option leading to group breakup. Previous attempts to synthesize these two models did not allow for the fact that the tug-of-war will affect group output, which in turn feeds back on the reproductive payments required by group members to remain cooperative. The bordered tug-of-war model, which does not assume complete reproductive control by any individual and allows for conflict within groups, predicts that the degree of within-group selfishness will increase as the noncooperative options become less attractive, e.g., as ecological constraints on solitary breeding increase. When the noncooperative option involves fighting for the group resource (e.g., territory) and leaving if the fight is lost, the subordinate's overall share of reproduction is predicted to be independent of its relatedness to the dominant and to increase the greater its probability of winning the fight, the less the value of the territory, and the greater its personal payoff for leaving. The unique predictions of the bordered tug-of-war model may fit skew data from a number of species, including meerkats, lions, and wood mice.