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Many animals rely on photoperiodic and non-photoperiodic environmental cues to gather information and appropriately time life-history stages across the annual cycle, such as reproduction, molt, and migration. Here, we experimentally demonstrate that the reproductive physiology, but not migratory behavior, of captive Pine Siskins (Spinus pinus) responds to both food and social cues during the spring migratory-breeding period. Pine Siskins are a nomadic finch with a highly flexible breeding schedule and, in the spring, free-living Pine Siskins can wander large geographic areas and opportunistically breed. To understand the importance of non-photoperiodic cues to the migratory-breeding transition, we maintained individually housed birds on either a standard or enriched diet in the presence of group-housed heterospecifics or conspecifics experiencing either the standard or enriched diet type. We measured body condition and reproductive development of all Pine Siskins and, among individually housed Pine Siskins, quantified nocturnal migratory restlessness. In group-housed birds, the enriched diet caused increases in body condition and, among females, promoted reproductive development. Among individually housed birds, female reproductive development differed between treatment groups, whereas male reproductive development did not. Specifically, individually housed females showed greater reproductive development when presented with conspecifics compared to heterospecifics. The highest rate of female reproductive development, however, was observed among individually housed females provided the enriched diet and maintained with group-housed conspecifics on an enriched diet. Changes in nocturnal migratory restlessness did not vary by treatment group or sex. By manipulating both the physical and social environment, this study demonstrates how multiple environmental cues can affect the timing of transitions between life-history stages with differential responses between sexes and between migratory and reproductive systems.

 

Our understanding of state-dependent behaviour is reliant on identifying physiological indicators of condition. Telomeres are of growing interest for understanding behaviour as they capture differences in biological state and residual lifespan. To understand the significance of variable telomere lengths for behaviour and test two hypotheses describing the relationship between telomeres and behaviour (i.e. the causation and the selective adoption hypotheses), we assessed if telomere lengths are longitudinally repeatable traits related to spring migratory behaviour in captive pine siskins ( Spinus pinus ). Pine siskins are nomadic songbirds that exhibit highly flexible, facultative migrations, including a period of spring nomadism. Captive individuals exhibit extensive variation in spring migratory restlessness and are an excellent system for mechanistic studies of migratory behaviour. Telomere lengths were found to be significantly repeatable ( R = 0.51) over four months, and shorter pre-migratory telomeres were associated with earlier and more intense expression of spring nocturnal migratory restlessness. Telomere dynamics did not vary with migratory behaviour. Our results describe the relationship between telomere length and migratory behaviour and provide support for the selective adoption hypothesis. More broadly, we provide a novel perspective on the significance of variable telomere lengths for animal behaviour and the timing of annual cycle events. 

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 the exploration of complex scenarios that present challenges to study in vivo but capture important complexity of the natural world.

 

The final stage of migration, when animals terminate migratory movements and transition to a more sedentary state, remains the least understood phase of migration. Whereas migrants that return to the same locations each year may use mechanisms associated with locating a specific destination, migrants with low site fidelity, such as nomadic migrants, may rely on local environmental cues to determine when to cease migratory movements. Using an experiment with captive birds, we tested whether the presence of a conspecific influences the termination of migration, indicated by changes in behaviour and physiology, in a nomadic migrant (the pine siskin, Spinus pinus ). We paired migratory birds with a non-migratory individual or left migratory and non-migratory individuals unpaired. Migratory paired birds had a significant decline in nocturnal activity immediately after pairing and activity levels remained lower two weeks later, with significant declines in energetic reserves and flight muscle size also observed. By contrast, migratory unpaired birds maintained high levels of activity and energetic reserves. These results provide evidence for a role of the social environment in migratory termination decisions. Social cues may be particularly useful in nomadic migrants, such as pine siskins, to facilitate settling in high quality, but unfamiliar, habitats. 

Many animals differentially express behaviours across the annual cycle as life stages are coordinated with seasonal environmental conditions. Understanding of the mechanistic basis of such seasonal changes in behaviour has traditionally focused on the role of changes in circulating hormone levels. However, it is increasingly apparent that other endocrine regulation mechanisms such as changes in local hormone synthesis and receptor abundance also play a role. Here I review what is known about seasonal changes in steroid hormone receptor abundance in relation to seasonal behaviour in vertebrates. I find that there is widespread, though not ubiquitous, seasonal variation in the expression of steroid hormone receptors in the brain, with such variation being best documented in association with courtship, mating and aggression. The most common pattern of seasonal variation is for there to be upregulation of sex steroid receptors with the expression of courtship and mating behaviours, when circulating hormone levels are also high. Less well-documented are cases in which seasonal increases in receptor expression could compensate for low circulating hormone levels or seasonal downregulation that could serve a protective function. I conclude by identifying important directions for future research. 

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.