When should signals of submission be given?-A game theory model.

During contests, losing animals often show signals communicating submission. From an evolutionary viewpoint, however, it is not so obvious why the losing individual gives such a signal instead of running away and why the winning individual accepts the signal instead of inflicting more severe damage. We investigated factors influencing the evolution of signals of submission using a numerical ESS model. The present analysis reveals that there is much space for the evolution of signals of submission, even when the winner of an escalated contest gets some extra benefit. In most cases, signals of submission are given by animals which are slightly or moderately weaker than the opponent. Signals of submission are expected to occur frequently (1) when the value of contested resource does not differ greatly from the cost of injury, (2) when the extra benefit of winning an escalated contest is small, (3) when the opportunity for safe retreat by the losing animal is small, and (4) when the estimation of the difference in the resource holding potential (RHP) between the combatants is accurate but not perfect.