Intensity of territorial defense in red squirrels: an experimental test of the asymmetric war of attrition

Summary Red squirrels defend exclusive, individual territories year round, 20% to 50% of females do not breed in any given year, and breeding females raise juveniles on their territories. Breeding is asynchronous, and the offspring of early-breeding females are more likely to hold an independently won territory than are late-born offspring. Based on the asymmetric war of attrition, we made the following predictions: (1) squirrels would respond more intensely to the calls of unfamiliar individuals than to the calls of neighbors; (2) breeding females would respond more intensely to unknown calls than would non-breeding females or males; (3) early-breeding females would respond more intensely than would late-breeding females to unknown calls; and (4) all classes of squirrels would respond similarly to the calls of neighbors. Playback experiments supported the predictions. Alternative hypotheses of kin selection, risk of infanticide, and seasonal difference in intruder pressure could not explain the results.