Competition for early arrival in migratory birds


Hanna Kokko, Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK. Fax: + 44 1223336676. E-mail:


1. It is widely accepted that the arrival order of migratory birds is correlated with the condition of the birds, which leads to high quality individuals occupying prime sites. However, the theoretical backgrounds for this argument have been lacking. A simple game-theoretic model of arrival timing is provided which investigates the evolutionary stability of condition-dependent arrival order in territorial migrant birds.

2. Competition for territories or other priority-dependent benefits can lead to arrival dates far preceding the cost-minimizing date (the optimum date in the absence of competition) for all but the weakest individuals. Increasing the number of competitors can generate a ‘cascading’ competition for early arrival, which advances arrival dates further apart from the individual optimum dates for the onset of breeding.

3. At equilibrium, arrival order corresponds strictly to condition order only if marginal costs of advancing arrival are always larger for individuals in lower condition. If spring mortality vacates territories for later-arriving birds, the criterion for ‘honest’ arrival order becomes still stricter: differential survival costs should exist, but survival differences among individuals (or, alternatively, territory quality differences) should not be very large.

4. If the habitat is saturated so that there is a risk of not obtaining a territory at all, or if worst territories are of much lower value than the rest, competition may lead to the majority of the population arriving within a fairly short interval, followed by a much later floating fraction. This synchrony in the arrival of breeders imposes an increasing cost for the lesser fit breeding birds. Thus, arrival costs paid are not necessarily highest for earliest arriving individuals, but for those who have the most to lose if they drop a few steps in the arrival order.

5. Competition for high quality territories can also lead to partial migration, in which case birds in good condition are expected to be most likely to remain resident.