This is a response to a recent article by Hanna Kokko and William J. Sutherland (American Naturalist 152:354-366), who consider evolutionarily stable territory acceptance rules for animals that face the decision between settling on a poor territory now (which is then retained for life) or waiting for better habitat to become available later (taking a chance of dying before reproducing). In contrast to these authors, we argue that the evolutionarily stable threshold quality above which territories are acceptable does depend on whether individuals compete for a single territory (queuing) or for multiple territories (floating) and also on whether access to territories is determined by a hierarchy among waiting individuals. More specifically, we show the following: First, if the choice is between floating and settling, the evolutionarily stable acceptance threshold is such that threshold territories yield an expected lifetime reproductive success (LRS) of [Formula: see text], the survival probability of a floater. Second, if the choice is between queuing and settling, the evolutionarily stable threshold may correspond to any LRS between [Formula: see text] and unity. Third, the number of nonbreeding individuals in the population is maximized at a threshold of unity. In other words, the evolutionarily stable threshold does not maximize the nonbreeding fraction of the population. We argue that models of territory choice should carefully specify the mechanism of choice because some choice processes (e.g., indiscriminate habitat use above the threshold) do not admit an evolutionarily stable acceptance rule.
In most species, some individuals delay reproduction or occupy inferior breeding positions. The queue hypothesis tries to explain both patterns by proposing that individuals strategically delay breeding (queue) to acquire better breeding or social positions. In 1995, Ens, Weissing, and Drent addressed evolutionarily stable queuing strategies in situations with habitat heterogeneity. However, their model did not consider the non-mutually exclusive individual quality hypothesis, which suggests that some individuals delay breeding or occupy inferior breeding positions because they are poor competitors.Here we extend their model with individual differences in competitive abilities, which are probably plentiful in nature. We show that including even the smallest competitive asymmetries will result in individuals using queuing strategies completely different from those in models that assume equal competitors. Subsequently, we investigate how well our models can explain settlement patterns in the wild, using a long-term study on oystercatchers. This long-lived shorebird exhibits strong variation in age of first reproduction and territory quality. We show that only models that include competitive asymmetries can explain why oystercatchers' settlement patterns depend on natal origin. We conclude that predictions from queuing models are very sensitive to assumptions about competitive asym-
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