The transition to a migratory state involves coordinated changes in physiology and behaviour. In species with regular, predictable (obligate) migrations, increasing day length triggers the expression of a spring migratory state and androgens play an important role in stimulating its development. By contrast, we know little about the environmental cues and endocrine mechanisms that regulate migration in species with less predictable (facultative) migrations. Here, we tested whether photoperiod stimulates a migratory state in a facultative nomadic migrant, the pine siskin (Spinus pinus). We exposed wintering birds to either a naturally increasing or short-day photoperiod and measured physiological and behavioural changes indicative of a migratory state. We also examined changes in circulating hormones that may play a role in the migratory transition. Natural-day, but not short-day, birds displayed physiological preparations for migration, including increases in fat deposition, and showed increased levels of migratory restlessness. We found no evidence for a role of corticosterone in the migratory transition, but testosterone may be important. This study is the first experimental test of the role of photoperiod in regulating facultative migration and demonstrates that the predictive cue used by many obligate migrants to time spring migration is also important in a facultative migrant.
Physiological preparations for migration generally reflect migratory strategy. Migrant birds fuel long-distance flight primarily with lipids, but carrying excess fuel is costly; thus, the amount of fat deposited prior to departure often reflects the anticipated flight duration or distance between refueling bouts. Seasonal pre-migratory deposition of fat is well documented in regular seasonal migrants, but is less described for more facultative species. We analyze fat deposits of free-living birds across several taxa of facultative migrants in the songbird subfamily Carduelinae, including house finches (Haemorhous mexicanus), American goldfinches (Spinus tristis), pine siskins (Spinus pinus) and four different North American ecotypes of red crossbills (Loxia curvirostra), to evaluate seasonal fat deposition during facultative migratory periods. Our data suggest that the extent of seasonal fat deposits corresponds with migratory tendency in these facultative taxa. Specifically, nomadic red crossbills with a seasonally predictable annual movement demonstrated relatively large seasonal fat deposits coincident with the migratory periods. In contrast, pine siskins, thought to be more variable in timing and initiation of nomadic movements, had smaller peaks in fat deposits during the migratory season, and the partial migrant American goldfinch and the resident house finch showed no peaks coincident with migratory periods. Within the red crossbills, those ecotypes that are closely associated with pine habitats showed larger peaks in fat deposits coincident with autumn migratory periods and had higher wing loading, whereas those ecotypes associated with spruces, Douglas-fir and hemlocks showed larger peaks coincident with spring migratory periods and lower wing loading. We conclude that population averages of fat deposits do reflect facultative migration strategies in these species, as well as the winter thermogenic challenges at the study locations. A difference in seasonal fattening and wing loading among red crossbill ecotypes is consistent with the possibility that they differ in their migratory biology, and we discuss these differences in light of crossbill reproductive schedules and phenologies of different conifer species.
Like many organisms, birds exhibit daily (circadian) and seasonal biological rhythms, and within populations both daily and seasonal timing often vary among individuals. Because photoperiod interacts with the circadian rhythms of many organisms to induce seasonal changes in behaviour and physiology, it is hypothesized that differences in daily timing, called chronotypes, underpin differences among individuals in the timing of seasonal events. For seasonal events stimulated by increasing daylength, this hypothesis predicts a positive relationship between the timing of daily and seasonal activities of individuals, with advanced chronotypes expressing events earlier in the year. The few previous tests of this hypothesis have focused on seasonal reproductive timing in birds. However, the hypothesis predicts that this relationship should extend to other photoinduced seasonal events. Therefore, we tested whether variation in chronotype was associated with variation in spring migratory timing in a captive songbird model, the pine siskin ( Spinus pinus ). We found that pine siskins expressing migratory restlessness exhibited repeatable chronotypes in their timing of nocturnal activity. Further, chronotype was significantly associated with the onset date of migratory behaviour, consistent with the hypothesized relationship between chronotype and seasonal timing.
The differential allocation hypothesis predicts individuals will increase their reproductive investment when mated to a high quality partner. In many species of fish with biparental care females prefer large males due to the males’ greater ability to raise more offspring to independence. I examined the relationship between mate quality, parental care and number of offspring in a natural population of convict cichlids Amatitlania siquia. The frequency of frontal displays by females was positively correlated with male standard length. Additionally, as males increased in length relative to their mate, females increased the frequency of chases towards predators, while males decreased the number of displays towards brood predators. This trade-off in parental effort within a pair due to mate quality is a key prediction of differential allocation. The number of offspring was correlated with male, but not female, standard length. These results support the differential allocation hypothesis in that females offered more parental care to offspring of a larger male, while their mates decreased the amount of care they provided. Additionally, females benefited in terms of number of offspring by pairing with higher quality mates. Increased female investment may provide an incentive to ensure male care and maintain pair bonding, which could lead to greater reproductive success through increased offspring survival.
Many organisms use environmental cues to time events in their annual cycle, such as reproduction and migration, with the appropriate timing of such events impacting survival and reproduction. As the climate changes, evolved mechanisms of cue use may facilitate or limit the capacity of organisms to adjust phenology accordingly, and organisms often integrate multiple cues to fine-tune the timing of annual events. Yet our understanding of how suites of cues are integrated to generate observed patterns of seasonal timing remains nascent. We present an overarching framework to describe variation in the process of cue integration in the context of seasonal timing. This framework incorporates both cue dependency and cue interaction. We then summarize how existing empirical findings across a range of vertebrate species and life cycle events fit into this framework. Finally, we use a theoretical model to explore how variation in modes of cue integration may impact the ability of organisms to adjust phenology adaptively in the face of climate change. Such a theoretical approach can facilitate exploration of complex scenarios that present challenges to study in vivo but capture important complexity of the natural world.
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