Environmental regulation of annual schedules in opportunistically-breeding songbirds: adaptive specializations or variations on a theme of white-crowned sparrow?

How birds use environmental cues to time breeding, migration and molt has been the subject of intensive study for nearly 90 years. Most work has focused on seasonal breeders; opportunistic breeders have been presumed to differ fundamentally from seasonal taxa in ways that facilitate coping with unpredictable environments. Understanding patterns and mechanisms of opportunists' responses to environmental cues can reveal the extent to which different environments require specialized adaptations of cue response systems. In this review we will present our perspective on how patterns and mechanisms of environmental cue response of three groups of opportunists--zebra finches, crossbills and Darwin's finches--compare with seasonal breeders. Long-standing predictions regarding tonic activity of the hypothalamic gonadotropin-releasing hormone system have been confirmed in at least some opportunists. However, opportunists resemble seasonal breeders in some surprising ways, illustrating basic similarity among taxa facing very different timing challenges. For instance, many opportunists completely regress the gonads outside breeding times, rely on initial predictive cues (both photic and non-photic) to regulate timing and rate of reproductive development, and in some cases even appear to display internal changes in responsiveness to environmental cues (i.e., cycles of reproductive refractoriness and sensitivity). Although advantages of unrestricted temporal flexibility are intuitively clear for animals coping with unpredictable habitats, the available data on these opportunists indicate that in all but the most extremely capricious situations the advantages of flexibility may be at least partly outweighed by contrasting advantages of following a reliable temporal schedule.