This paper advances an hypothesis that the primary adaptive driver of seasonal migration is maintenance of site fidelity to familiar breeding locations. We argue that seasonal migration is therefore principally an adaptation for geographic persistence when confronted with seasonality -analogous to hibernation, freeze tolerance, or other organismal adaptations to cyclically fluctuating environments. These ideas stand in contrast to traditional views that bird migration evolved as an adaptive dispersal strategy for exploiting new breeding areas and avoiding competitors. Our synthesis is supported by a large body of research on avian breeding biology that demonstrates the reproductive benefits of breeding-site fidelity. Conceptualizing migration as an adaptation for persistence places new emphasis on understanding the evolutionary trade-offs between migratory behaviour and other adaptations to fluctuating environments both within and across species. Seasonality-induced departures from breeding areas, coupled with the reproductive benefits of maintaining breeding-site fidelity, also provide a mechanism for explaining the evolution of migration that is agnostic to the geographic origin of migratory lineages (i.e. temperate or tropical). Thus, our framework reconciles much of the conflict in previous research on the historical biogeography of migratory species. Although migratory behaviour and geographic range change fluidly and rapidly in many populations, we argue that the loss of plasticity for migration via canalization is an overlooked aspect of the evolutionary dynamics of migration and helps explain the idiosyncratic distributions and migratory routes of long-distance migrants. Our synthesis, which revolves around the insight that migratory organisms travel long distances simply to stay in the same place, provides a necessary evolutionary context for understanding historical biogeographic patterns in migratory lineages as well as the ecological dynamics of migratory connectivity between breeding and non-breeding locations.

Seasonal migration is intrinsically connected to the balance of survival and reproduction, but whether migratory behavior influences species’ position on the slow-fast continuum of life history is poorly understood. We found that boreal-breeding birds that migrate long distances exhibit higher annual adult survival and lower annual reproductive investment relative to co-distributed boreal species that migrate shorter distances to winter closer to their breeding grounds. Our study uses “vital rates” data on reproductive output and survivorship compiled from the literature for a species assemblage of 45 species of mostly passerine birds. These species breed sympatrically in North American boreal forests but migrate to a diversity of environments for the northern winter. After controlling for body size and phylogeny, migration distance and apparent annual adult survival are positively related across species. Both migration distance and survival are positively correlated with wintering in environments that are warmer, wetter, and greener. At the same time, longer migrations are associated with reduced time spent on the breeding grounds, lower clutch sizes, and lower fecundity (clutch size × maximum number of broods per year). Although seasonal migration is often associated with high mortality, our results suggest that long-distance migration imposes selection pressures that both confer and demand high adult survival rates. That is, owing to the reproductive cost of long-distance migration, this strategy can only persist if balanced by high adult survival. Our study supports the idea that migration evolves to promote survival of species breeding in seasonal environments. In boreal birds, the evolution of the longest migrations yields the highest survival, but at an inherent cost to annual fecundity. Our results therefore reveal migratory distance as a fundamental axis of the slow-fast continuum that predicts, and is inextricable from, the balance of survival and reproduction.

Animals must balance various costs and benefits when deciding when to breed. The costs and benefits of breeding at different times have received much attention, but most studies have been limited to investigating short‐term season‐to‐season fitness effects. However, breeding early, versus late, in a season may influence lifetime fitness over many years, trading off in complex ways across the breeder’s lifespan. In this study, we examined the complete life histories of 867 female tree swallows (Tachycineta bicolor) breeding in Ithaca, New York, between 2002 and 2016. Earlier breeders outperformed later breeders in short‐term measures of reproductive output and offspring quality. Though there were weak indications that females paid long‐term future survival costs for breeding early, lifetime fledgling output was markedly higher overall in early‐breeding birds. Importantly, older females breeding later in the season did not experience compensating life history advantages that suggested an alternative equal‐fitness breeding strategy. Rather, most or all of the swallows appear to be breeding as early as they can, and differences in lay dates appear to be determined primarily by differences in individual quality or condition. Lay date had a significant repeatability across breeding attempts by the same female, and the first lay date of females fledged in our population was strongly influenced by the first lay date of their mothers, indicating the potential for ongoing selection on lay date. By examining performance over the entire lifespan of a large number of individuals, we were able to clarify the relationship between timing of breeding and fitness and gain new insight into the sources of variability in this important life history trait.

Divergence in song between allopatric populations can contribute to premating reproductive isolation in territorial birds. Song divergence is typically measured by quantifying divergence in vocal traits using audio recordings, but field playback experiments provide a more direct way to behaviorally measure song divergence between allopatric populations. The White-breasted Wood-Wren (Henicorhina leucosticta; hereafter “WBWW”) is an abundant Neotropical species with four mitochondrial clades (in Central America, the Darién, the Chocó and the Amazon) that are deeply divergent (~5–16% sequence divergence). We assessed the possibility that the WBWW as currently defined may represent multiple biological species by conducting both statistical analysis of vocal characters and field playback experiments within three clades (Central America, Chocó and Amazon). Our analysis of vocal traits revealed that Central American songs overlapped in acoustic space with Chocó songs, indicating vocal similarity between these two populations, but that Central American songs were largely divergent from Amazonian songs. Playback experiments in the Caribbean lowlands of Costa Rica revealed that Central American WBWWs typically responded aggressively to songs from the Chocó population but did not respond to playback of songs from the Amazonian population, echoing the results of the vocal trait analysis. This marked difference in behavioral response demonstrates that the songs of Central American and Amazonian WBWWs (but not Central American and Chocó WBWWs) have diverged sufficiently that Central American WBWWs no longer recognize song from Amazonian WBWWs as a signal to elicit territorial defense. This suggests that significant premating reproductive isolation has evolved between these two populations, at least from the perspective of the Central American population, and is consistent with the possibility that Central American and Amazonian populations represent distinct biological species. We conclude by advocating for the further use of field playback experiments to assess premating reproductive isolation (and species limits) between allopatric songbird populations, a situation where behavioral systematics can answer questions that phylogenetic systematics cannot.

What is already known: Experimental manipulations of glucocorticoids during development affect various phenotypic traits, including morphology, HPA axis functioning, and behavior.Most such studies have manipulated glucocorticoids to mimic an increase in baseline glucocorticoid levels, rather than an acute stress response. Because most manipulations have been done in captive populations, impacts of glucocorticoids on phenotype, movement patterns, and fitness under natural conditions are not well understood.What this study adds: Using a wild, free-living songbird, we demonstrate that just five brief, acute increases in the glucocorticoid stress hormone corticosterone during development are sufficient to produce phenotypic effects including reduced mass, higher baseline and stressinduced corticosterone levels, and shorter telomeres. We did not find evidence that this manipulation affected time to maturity, post-fledging behavior, or survival. AbstractOrganisms are frequently exposed to challenges during development, such as poor weather and food shortage. Such challenges can initiate the hormonal stress response, which involves secretion of glucocorticoids. Although the hormonal stress response helps organisms deal with challenges, long-term exposure to high levels of glucocorticoids can have morphological, behavioral, and physiological consequences, especially during development. Glucocorticoids are also associated with reduced survival and telomere shortening. To investigate whether brief, acute exposures to glucocorticoids can also produce these phenotypic effects in free-living birds, we exposed wild tree swallow (Tachycineta bicolor) nestlings to a brief exogenous dose of cort once per day for five days and then measured their morphology, baseline and stress-induced corticosterone levels, and telomere length. We also deployed radio tags on a subset of nestlings, which allowed us to determine the age at which tagged nestlings left the nest (fledged) and their pattern of presence and absence at the natal site during the post-breeding period. Corticosteronetreated nestlings had lower mass, higher baseline and stress-induced corticosterone, and reduced telomeres; other metrics of morphology were affected weakly or not at all. Our treatment resulted in no significant effect on survival to fledging, fledge age, or age at first departure from the natal site, and we found no negative effect of corticosterone on inter-annual return rate.These results show that brief acute corticosterone exposure during development can have measurable effects on phenotype in free-living tree swallows. Corticosterone may therefore mediate correlations between rearing environment and phenotype in developing organisms, even in the absence of prolonged stressors.