The evolution of aggressive losers

We examine the question of when aggressive behavior of likely losers should be part of an evolutionarily stable strategy. We modified an earlier model by the authors that found situations where likely losers initiate aggressive interactions more often than likely winners. The modifications allowed us to examine the robustness of the previous study by including an unusually high number of possible strategies (n = 81) and to examine a wide range of parameter settings. First, we show that restricting attention to only a few most plausible strategies may change the overall results. Second, within the space where escalation is predicted, for a large percentage of the parameter settings (85%), an ESS exists that leads to a somewhat counterintuitive situation where escalation is more often initiated by the likely loser than by the likely winner of the contest. In contrast, an ESS that favors escalation by likely winners was found only for about 3% of parameter settings. Furthermore, we use simulations of evolution in a finite population to verify for certain parameter settings that the analytically predicted ESS's could in fact evolve. Our results suggest that ESSs in which the likely loser rather than the likely winner is expected to initiate escalation are generic and ESSs in which the opposite is true need to be explained by incorporating specific features of the biology of a given species into more detailed models.