Social environment influences termination of nomadic migration.

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.

Declining food availability, corticosterone, and migratory response in a nomadic, irruptive migrant.

While obligate migrants time their movements to respond to predictable changes in the environment, facultative migration is characterized by more variable movements that are driven by unpredictable changes in resource availability. The proximate cues that trigger facultative migrations and the endocrine mechanisms involved in these responses remain poorly understood, though corticosterone may be a key mediator of facultative migration due to its effects on activity and metabolic processes. We conducted experiments in the fall and spring to examine the response of pine siskins (Spinus pinus), a facultative migrant, to a two-stage food restriction. Our goals were to examine whether declining food availability stimulated behavioral and/or physiological changes consistent with a migratory response, whether anticipatory behavioral and physiological adjustments occurred when birds were initially presented with changing food availability, and if observed changes corresponded to changes in circulating corticosterone levels. We found no evidence of preparatory physiological changes for migration, but food-restricted birds in the spring had increased daytime activity indicative of a migratory response. Corticosterone increased at each stage of the restriction and the change in corticosterone corresponded to the magnitude of decline in body condition. Increased corticosterone was also correlated with larger increases in activity during the initial stage of food restriction, but only during the spring, when birds also displayed higher levels of initial body condition. These results suggest that initial energetic state and corticosterone response may interact to determine an individual's behavioral and physiological response to declining food availability and ultimately the facultative migratory response.

Variation in chronotype is associated with migratory timing in a songbird.

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.

Environmental Cue Integration and Phenology in a Changing World.

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.

Increasing photoperiod stimulates the initiation of spring migratory behaviour and physiology in a facultative migrant, the pine siskin.

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.