Developmental temperature predicts the adult response to stressors in a free-living passerine.

Early-life conditions can have substantial effects on the ways animals respond to stressors as adults. In particular, thermal conditions during development affect juveniles' responses to stressors, and there is evidence that these effects may extend into adulthood. However, these effects remain poorly understood, especially in free-living organisms. We test the prediction that ambient temperatures during laying, embryonic development and nestling development affect the hormonal mediators of the response to stressors in adults. To do so, we use a long-term dataset of tree swallows (Tachycineta bicolor) with records from both natal development and adult breeding. We found a strong, negative relationship between ambient temperature during early development (incubation) and an individual's corticosterone (CORT) response to stress later in life (while incubating her own young). Thermal conditions during other stages of natal development also showed a weak relationship with baseline CORT during provisioning. In a post hoc analysis, we found no evidence that ambient temperature during development differentially influenced the survival and recruitment of juveniles with different CORT phenotypes. Our results show that thermal conditions during development can have long-term effects on how individuals respond to stressors.

Does migration constrain glucocorticoid phenotypes? Testing corticosterone levels during breeding in migratory versus resident birds.

Corticosterone, the main glucocorticoid in birds, is a major mediator of the incredible physiological feat of migration. Corticosterone plays important roles in migration, from preparation to in-flight energy mobilization to refueling, and corticosterone levels often show distinct elevations or depressions during certain stages of the migratory process. Here, we ask whether corticosterone's role in migration shapes its modulation during other life history stages, as is the case with some other phenotypically flexible traits involved in migration. Specifically, we use a global dataset of corticosterone measures to test whether birds' migratory status (migrant versus resident) predicts corticosterone levels during breeding. Our results indicate that migratory status predicts neither baseline nor stress-induced corticosterone levels in breeding birds; despite corticosterone's role in migration, we find no evidence that migratory corticosterone phenotypes carry over to breeding. We encourage future studies to continue to explore corticosterone in migrants versus residents across the annual cycle. Additionally, future efforts should aim to disentangle the possible effects of environmental conditions and migratory status on corticosterone phenotypes; potentially fruitful avenues include focusing on regions where migrants and residents overlap during breeding. Overall, insights from work in this area could demonstrate whether migration shapes traits during other important life stages, identify tradeoffs or limitations associated with the migratory lifestyle, and ultimately shed light on the evolution of flexible traits and migration.

Developmental stage-dependent effects of perceived predation risk on nestling tree swallows (Tachycineta bicolor).

The risk of predation directly affects the physiology, behavior, and fitness of wild birds. Strong social connections with conspecifics could help individuals recover from a stressful experience such as a predation event; however, competitive interactions also have the potential to exacerbate stress. Few studies have investigated the interaction between environmental stressors and the social landscape in wild bird populations. In 2 years of field studies, we experimentally simulated predation attempts on breeding female tree swallows (Tachicyneta bicolor). At the same time, we manipulated female breast plumage color, a key social signal. Simulated predation events on tree swallows early in the nestling period reduced young nestlings' mass by approximately 20% and shortened telomere lengths. Ultimately, only 31% of nestlings in the predation group fledged compared with 70% of control nestlings. However, the effects of experimental manipulations were timing dependent: the following year when we swapped the order of the experimental manipulations and simulated predation during incubation, there were no significant effects of predation on nestling condition or fledging success. Contrary to our expectations, manipulation of the social environment did not affect the response of tree swallows to simulated predation. However, manipulating female plumage during the nestling period did reduce nestling skeletal size and mass, although the effects depended on original plumage brightness. Our data demonstrate that transient stressors on female birds can have carry-over effects on their nestlings if they occur during critical periods in the breeding season.

Social signal manipulation and environmental challenges have independent effects on physiology, internal microbiome, and reproductive performance in tree swallows (Tachycineta bicolor).

The social environment that individuals experience appears to be a particularly salient mediator of stress resilience, as the nature and valence of social interactions are often related to subsequent health, physiology, microbiota, and overall stress resilience. Relatively few studies have simultaneously manipulated the social environment and ecological challenges under natural conditions. Here, we report the results of experiments in wild tree swallows (Tachycineta bicolor) in which we manipulated both ecological challenges (predator encounters and flight efficiency reduction) and social interactions (by experimental dulling of a social signal). In two experiments conducted in separate years, we reversed the order of these treatments so that females experienced either an altered social signal followed by a challenge or vice-versa. Before, during, and after treatments were applied, we tracked breeding success, morphology and physiology (mass, corticosterone, and glucose), nest box visits via an RFID sensor network, cloacal microbiome diversity, and fledging success. Overall, we found that predator exposure during the nestling period reduced the likelihood of fledging and that signal manipulation sometimes altered nest box visitation patterns, but little evidence that the two categories of treatment interacted with each other. We discuss the implications of our results for understanding what types of challenges and what conditions are most likely to result in interactions between the social environment and ecological challenges.

A virtual bird's eye view: Live streaming nest boxes to continue outreach in the era of COVID-19.

COVID-19 created a host of challenges for science education; in our case, the pandemic halted our in-person elementary school outreach project on bird biology. This project was designed as a year-long program to teach fifth-grade students in Ithaca, New York, USA, about bird ecology and biodiversity using in-person presentations, games, activities, and outdoor demonstrations. As a central part of this effort, we set up nest boxes on school property and planned to monitor them with students during bird breeding in the spring. Here, we describe our experiences transitioning this program online: we live streamed nest boxes to the students' virtual classroom and used them as a focal point for virtual lessons on bird breeding and nestling development. In an era of social distancing and isolation, we propose that nest box live streaming and virtual lessons can support communities by providing access to the outdoors and unconventional science learning opportunities for all students. Instituting similar programs at local schools has the potential to increase equitable learning opportunities for students across geographic locations and with varying degrees of physical access to the outdoors and nature.

Effects of Artificial Light at Night on Avian Provisioning, Corticosterone, and Reproductive Success.

Artificial light at night (hereafter "ALAN") affects 88% of the land area in Europe and almost half of the land area in the USA, with even rural areas exposed to lights from agricultural and industrial buildings. To date, there have been few studies that assess the impacts of ALAN on both wildlife behavior and physiology. However, ALAN may alter energy expenditure and/or stress physiology during the breeding period, potentially reducing reproductive success and resulting in conservation implications. Here, we experimentally exposed adult female and nestling tree swallows (Tachycineta bicolor) to ALAN. We then measured the effects of ALAN compared with control conditions on parental behavior (provisioning rate), nestling physiology (corticosterone levels), and reproductive success (likelihood of all eggs hatching and all nestlings fledging per nest). Our results showed that ALAN-exposed females provisioned their nestlings at lower rates than control females. Although relatively weak, our results also suggested that ALAN-exposed nestlings had reduced baseline and increased stress-induced corticosterone compared with control nestlings. ALAN-exposed nestlings also showed greater negative feedback of circulating corticosterone. We found no support for our prediction that ALAN would reduce nestling body condition. Finally, we found some support for a negative effect of ALAN on the likelihood that all eggs hatched in a given nest, but not the likelihood that all nestlings fledged. Therefore, while it is possible that the behavioral and physiological changes found here result in long-term consequences, our results also suggest that direct ALAN exposure alone may not have substantially large or negative effects on tree swallows. Exposure regimes for free-living birds, such as exposure to a combination of anthropogenic disturbances (i.e., ALAN and noise pollution) or direct and indirect effects of ALAN (i.e., effects on physiology due to direct light exposure and alterations in food availability), may produce different results than those found here.

Full lifetime perspectives on the costs and benefits of lay-date variation in tree swallows.

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.

HormoneBase, a population-level database of steroid hormone levels across vertebrates.

Hormones are central regulators of organismal function and flexibility that mediate a diversity of phenotypic traits from early development through senescence. Yet despite these important roles, basic questions about how and why hormone systems vary within and across species remain unanswered. Here we describe HormoneBase, a database of circulating steroid hormone levels and their variation across vertebrates. This database aims to provide all available data on the mean, variation, and range of plasma glucocorticoids (both baseline and stress-induced) and androgens in free-living and un-manipulated adult vertebrates. HormoneBase (www.HormoneBase.org) currently includes >6,580 entries from 476 species, reported in 648 publications from 1967 to 2015, and unpublished datasets. Entries are associated with data on the species and population, sex, year and month of study, geographic coordinates, life history stage, method and latency of hormone sampling, and analysis technique. This novel resource could be used for analyses of the function and evolution of hormone systems, and the relationships between hormonal variation and a variety of processes including phenotypic variation, fitness, and species distributions.