Modelling migration in birds: competition's role in maintaining individual variation.

Animals exhibit extensive intraspecific variation in behaviour. Causes of such variation are less well understood. Here, we ask when competition leads to the maintenance of multiple behavioural strategies. We model variability using the timing of bird migration as an example. Birds often vary in when they return from non-breeding grounds to establish breeding territories. We assume that early-arriving birds (counting permanent residents as 'earliest') select the best territories. But arriving before the optimal (frequency-independent) breeding date incurs a fitness penalty. Using simulations, we find stable sets of return dates. When year-round residency is viable, the greatest between-individual variation occurs when a small proportion of permanent residents is favoured, and the rest of the population varies in their return times. However, when fitness losses due to year-round residency exceed the benefits of breeding in the worst territory, all individuals migrate, although their return dates often vary continuously. In that case, individual variation is inversely related to fitness risks and positively related to territory inequality. This result is applicable across many systems: when there is more to gain through competition, or when its risks are small, a diversity of individual strategies prevails. Additionally, stability can depend upon the distribution of resources.