Emergence of games from ecological trade-offs: longevity changes strategies for extra-pair mating in birds

Abstract

Each member of a breeding pair benefits if the other does more of the parental investment, so there is scope for behaviours that can be interpreted as both cooperative and competitive games between males and females. Extra-pair mating, widespread among socially monogamous birds, adds extra conflict but also potential opportunity to these social interactions. We analyse an individual-based model of a social environment with simple behavioural strategies where game-like patterns and cooperative outcomes emerge. The model focuses on three evolving traits: female propensity for extra-pair copulations and male investment in territorial behaviour and care. Male traits are reaction norms that use experienced within and extra-pair copulations as information input. We found that female extra-pair mating provided incentives for males to reduce territorial aggression and increase care for offspring. However, when adult survival was higher, male investment in care and territoriality changed from being negatively to positively correlated. This happened because longer life expectancy gave more behavioural opportunities for males, where nest desertion maximises lifetime male fitness when female extra-pair copulation is high. This outcome evolved gradually, with stable periods of intermediate extra-pair mating and low territoriality. These were punctuated by cycles of high extra-pair mating, nest desertion, reduced extra-pair mating and relapse to aggressive territoriality before a new stable phase was established. Each successive trait cycle was faster and smaller, indicating that through evolution of reaction norms, the gene pool has a long history that canalizes the evolution of behaviours, which can be interpreted as emergence and refinement of frequency-dependent games.Significance statementMost birds that mate in monogamous pairs engage in extra-pair copulations. Males of some species invest (in varying proportions) in offspring that are genetically related only to their social female. The great variability of extra-pair mating levels among different species and populations, supported by numerous field studies, indicates that social and ecological factors play a crucial role in shaping the behaviour. We analyse a computer model in which extra-pair mating evolves concurrently with male reproductive investments in territory defence and offspring care and show how common ecological trade-offs may change the social dynamics within a breeding population and lead to the emergence of complex social interactions between males, females and their neighbours.