Population density and the evolution of male aggression

In some cases male animals engage in aggressive contests for access to females, in others they adopt more passive strategies and invest in traits that assist them in detecting females or in competing with rivals in other ways, such as sperm competition. One possible factor determining the fitness of these different strategies is population density. Theoretically, aggressive tactics should be found at intermediate population densities. At low densities males that invest in traits related to searching for mates could be favoured, whereas at the highest densities males that fight over females might pay excessive costs for this behaviour because of the number of rival males that they will encounter. Current empirical evidence is mostly consistent with this scheme: in some cases it seems that traits that are associated with locating mates are favoured at low densities, with aggression related traits favoured at higher densities, and in other cases aggression is selected but as density increases less aggressive strategies become more common. There remain substantial differences between species, however, and I discuss how variation in mating system, in the costs of aggression and in the nature of sperm competition, plus ecological differences between species, can change the relationship between population density and the fitness consequences of aggressive and passive behavioural strategies.