Long-distance migratory shorebirds travel faster towards their breeding grounds, but fly faster post-breeding.

Long-distance migrants are assumed to be more time-limited during the pre-breeding season compared to the post-breeding season. Although breeding-related time constraints may be absent post-breeding, additional factors such as predation risk could lead to time constraints that were previously underestimated. By using an automated radio telemetry system, we compared pre- and post-breeding movements of long-distance migrant shorebirds on a continent-wide scale. From 2014 to 2016, we deployed radio transmitters on 1,937 individuals of 4 shorebird species at 13 sites distributed across North America. Following theoretical predictions, all species migrated faster during the pre-breeding season, compared to the post-breeding season. These differences in migration speed between seasons were attributable primarily to longer stopover durations in the post-breeding season. In contrast, and counter to our expectations, all species had higher airspeeds during the post-breeding season, even after accounting for seasonal differences in wind. Arriving at the breeding grounds in good body condition is beneficial for survival and reproductive success and this energetic constraint might explain why airspeeds are not maximised in the pre-breeding season. We show that the higher airspeeds in the post-breeding season precede a wave of avian predators, which could suggest that migrant shorebirds show predation-minimizing behaviour during the post-breeding season. Our results reaffirm the important role of time constraints during northward migration and suggest that both energy and predation-risk constrain migratory behaviour during the post-breeding season.

Color expression in experimentally regrown feathers of an overwintering migratory bird: implications for signaling and seasonal interactions.

Plumage coloration in birds plays a critical role in communication and can be under selection throughout the annual cycle as a sexual and social signal. However, for migratory birds, little is known about the acquisition and maintenance of colorful plumage during the nonbreeding period. Winter habitat could influence the quality of colorful plumage, ultimately carrying over to influence sexual selection and social interactions during the breeding period. In addition to the annual growth of colorful feathers, feather loss from agonistic interactions or predator avoidance could require birds to replace colorful feathers in winter or experience plumage degradation. We hypothesized that conditions on the wintering grounds of migratory birds influence the quality of colorful plumage. We predicted that the quality of American redstart (Setophaga ruticilla) tail feathers regrown after experimental removal in Jamaica, West Indies, would be positively associated with habitat quality, body condition, and testosterone. Both yearling (SY) and adult (ASY) males regrew feathers with lower red chroma, suggesting reduced carotenoid content. While we did not observe a change in hue in ASY males, SY males shifted from yellow to orange plumage resembling experimentally regrown ASY feathers. We did not observe any effects of habitat, testosterone, or mass change. Our results demonstrate that redstarts are limited in their ability to adequately replace colorful plumage, regardless of habitat, in winter. Thus, feather loss on the nonbreeding grounds can affect social signals, potentially negatively carrying over to the breeding period.

Experimental and observational studies of seasonal interactions between overlapping life history stages in a migratory bird.

Prior to reproduction, migratory animals are at the juxtaposition of three life history stages in which they must finish the non-breeding stage, initiate and complete migration, and prepare for the onset of breeding. However, how these stages interact with one another is not fully understood. We provide evidence that, for migratory birds that begin breeding development prior to departure from non-breeding sites, the level of breeding preparation can drive migration phenology, a critical behavioral determinant of reproductive success. Specifically, male American redstart (Setophaga ruticilla) plasma androgen levels, which increase in males during the period leading into migration, were positively correlated with energetic condition. We empirically tested the hypothesis that elevated androgen simultaneously supports migratory and breeding preparation in a hormone manipulation field experiment. Males with testosterone implants showed advanced preparation for migration and breeding, and ultimately departed on migration earlier than controls. It is assumed that early departure leads to early arrival at breeding areas, which increases breeding success. Collectively, our observational and experimental results demonstrate how overlapping life history stages can interact to influence important components of an individual's fitness. This highlights the critical need for understanding population processes across the full life cycle of an organism to better understand the ecological and evolutionary origins of complex life history events.

Early elevation of testosterone advances migratory preparation in a songbird.

The timing of events associated with spring migration can be an important determinant of fitness for migratory birds. The need to prepare and maintain physical condition for migration is one demand that must be met for early arrival on breeding areas, and this demand is compounded by the energetic demands of the physiological transition to breeding. Here, we examined whether migratory birds, specifically males, can adequately meet both of these demands by elevating the sex steroid testosterone early during migratory preparation. To test this, we used a captive experiment in which we photostimulated male dark-eyed juncos (Junco hyemalis) and manipulated circulating testosterone. We predicted males with elevated testosterone would prepare to migrate and breed faster than control males or males where the ability of testosterone to bind with receptors, or be converted to estradiol, was inhibited (testosterone inhibited). We measured migratory preparation using mass, food intake, fat deposits and nocturnal activity (Zugunruhe). To estimate breeding preparation, we measured the diameter of the cloacal protuberance (CP). We found that males in the testosterone treatment group began migratory preparation approximately 10 days before controls. There was no difference in the magnitude of peak migratory condition between testosterone-implanted birds and controls. Males implanted with testosterone also reached the onset of breeding preparation faster and had larger peak CP diameter compared with controls. Testosterone-inhibited birds showed no signs of migratory preparation and only a weak increase in CP diameter. These results demonstrate that early elevation of testosterone during migratory preparation could incur benefits for males in terms of the ability to depart earlier from non-breeding areas and arrive in a more advanced breeding condition. This experiment demonstrates that there may be important physiological underpinnings to known winter to breeding season carry-over effects in migratory birds.

Corticosterone manipulations alter morph-specific nestling provisioning behavior in male white-throated sparrows, Zonotrichia albicollis.

In the polymorphic white-throated sparrow (Zonotrichia albicollis), tan-striped males provision nestlings at higher rates than do white-striped males. In a previous study, we found that tan-striped males had lower baseline corticosterone levels than white-striped males during the nestling stage. To determine if this variation in corticosterone influences morph-specific differences in nestling provisioning behavior, we used intraperitoneal osmotic pumps to increase baseline corticosterone levels in tan-striped males (TS CORT) and administer RU486, a glucocorticoid receptor antagonist, in white-striped males (WS RU486). These manipulations essentially reversed morph-specific nestling provisioning behavior in males. TS CORT males fed nestlings at lower rates than TS controls (vehicle-only implant), and at similar rates to WS controls (vehicle-only implant), while WS RU486 males fed nestlings at higher rates than WS controls, and at similar rates to TS controls. These results demonstrate that (1) increases in baseline corticosterone (i.e., below concentrations associated with the adrenocortical response to stress) can directly or indirectly inhibit nestling provisioning behavior, and (2) corticosterone influences morph-specific variation in parental behavior in male white-throated sparrows. This study contributes to the growing evidence that modulating baseline CORT mediates parental care and self-maintenance activities in birds, and thus may serve as a mechanism for balancing current reproductive success with survival.

Does stress response predict return rate in a migratory bird species? A study of American redstarts and their non-breeding habitat.

In vertebrates, the adrenocortical stress response activates an emergency life-history stage, which is thought to promote survival by helping individuals escape life-threatening situations. Although the adrenocortical stress response promotes many behavioural and physiological changes, it remains unclear whether this stress response actually translates into higher survival in wild vertebrates. We measured the adrenocortical stress response of non-breeding American redstarts (Setophaga ruticilla), a migratory bird that wintered in habitats of either high (mangroves) or low suitability (scrubs), and subsequently monitored their return rate during the following non-breeding seasons. The intensity of the adrenocortical stress response was consistent within individuals across the non-breeding season and was positively correlated with return rates in redstarts that wintered in scrubs, but not in redstarts that wintered in mangroves. Thus, in a context-dependent manner, the ability of an individual to physiologically react to stress determines its ability of returning to its non-breeding territory the following winters. For an individual, the ability to mount an important adrenocortical stress response probably benefits to survival. However, this beneficial effect probably depends on an individual's environment and phenotypic characteristics because these two variables are likely to affect its probability of being confronted with life-threatening stressors during its annual life cycle.

Circulating prolactin and corticosterone concentrations during the development of migratory condition in the Dark-eyed Junco, Junco hyemalis.

Endogenous plasma prolactin and baseline corticosterone concentrations were measured in Dark-eyed Juncos (Junco hyemalis, n=27) photostimulated into migratory condition to look at how these hormones may be linked to the development of migratory condition. In addition to the commonly used assay for corticosterone, a recombinant-derived European starling prolactin assay validated for Dark-eyed juncos was used to measure endogenous prolactin in order to detect small but significant changes in plasma prolactin levels. In response to transfer from short (10.5L:13.5D) to long (18L:6D) days, the birds increased in body mass, fat score, daily food intake, and nocturnal migratory locomotor activity (Zugunruhe). On short-days, both hormones were low (corticosterone mean=2.89ng/mL+/-0.48 SE; prolactin mean=6.43ng/mL+/-1.31 SE). But, within 14 days of photostimulation both hormones increased significantly (Day 14: corticosterone mean=5.71ng/mL+/-0.77 SE; prolactin mean=19.67ng/mL+/-2.81 SE), rising further by Day 48 (corticosterone mean=8.41ng/mL+/-0.72; prolactin mean=112.67ng/mL+/-9.18 SE). On Day 48, birds with the most fat (fat score=3) had significantly higher corticosterone levels than those with less fat (fat score=2). This pattern, albeit not statistically significant, was similar for prolactin. These results illustrate that, independent of the seasonal peak in prolactin associated with the onset of photorefractoriness, plasma prolactin levels can rise, in concert with corticosterone, as birds come into spring migratory condition, providing some support for earlier hypotheses that these two hormones play an integral role in the development of migratory condition. Whether similar changes in plasma prolactin occur with respect to autumn migration, as does baseline corticosterone, has yet to be determined.

Intraperitoneal delivery of exogenous corticosterone via osmotic pump in a passerine bird.

Selecting the most reliable method to deliver exogenous steroids remains a problem for researchers, particularly when designing experiments on small birds. We used intraperitoneal (IP) osmotic pumps to deliver exogenous corticosterone (CORT) and RU486 in captive White-throated Sparrows. Males received implants containing either a low (LD) or moderate dose (MD) of CORT, RU486 (RU), or polyethylene glycol vehicle only (V). This method provided sustained elevations in baseline CORT in both LD and MD males compared to V males, with higher CORT levels induced in MD males. Slight increases in post-implant CORT resulted in V males, but not in RU males. We observed no significant change in the condition of V males in terms of body mass, furcular fat score or food intake rates, although locomotor activity declined slightly after implantation. Taken together, our results suggest that IP pumps had little impact on the overall health of the birds. CORT levels were maintained within natural ranges for this species, suggesting that our results are biologically relevant and useful for future endocrine studies with small birds.

Endocrine and behavioral response to a decline in habitat quality: effects of pond drying on the slider turtle, Trachemys scripta.

The effect of the simulated drying of a pond on the behavior and corticosterone secretion of Trachemys scripta was measured in a field situation. Slider turtles were held in experimental and control ponds (12 x 15 m) enclosed with a drift fence integrated with spring-triggered livetraps. The experimental pond water level was dropped 10 cm per day for 8 d, until water was completely drained. Slider turtles responded to the draining of the pond by the emigration of the majority (75%) of the experimental population. Emigrating turtles had significantly elevated corticosterone at Time 0 (blood sample within 10 min of handling=4.48 ng/mL+/-0.503SE) when compared with turtles captured in a control pond (Time 0=0.954 ng/mL+/-0.121SE), where conditions were held constant. Turtles emigrated during the final 72 hr of pond draining when ponds reached 30 cm depth and lower and water temperature was at least 30.8 degrees C or higher. Additionally, the effect of trapping using spring-activated livetraps was tested. Turtles held in livetraps (n=6) for 45-110 min showed a characteristic corticosterone response (Time 0=0.957 ng/mL+/-0.091SE; Time 30=2.85 ng/mL+/-0.131SE), indicating that this trapping technique alone does not stimulate corticosterone secretion. The findings of the study met our predictions that turtles would respond to the draining of the pond behaviorally by emigrating from the habitat concurrent with an elevated corticosterone concentration. This supports the view that corticosterone is involved in stress avoidance mechanisms that allow organisms to respond to environmental perturbations.

Corticosterone levels as indicators of habitat quality: effects of habitat segregation in a migratory bird during the non-breeding season.

During the non-breeding season, many species of territorial migratory birds exhibit a non-random pattern of habitat distribution, with males and females occupying different habitats. In this study, we examined possible physiological consequences arising from such habitat segregation in one migrant passerine species, the American redstart (Setophaga ruticilla), on its non-breeding grounds in Jamaica, West Indies. For 2 years, we measured concentrations of corticosterone, at the time of capture (baseline) and 30 min after capture (profile of acute corticosterone secretion), in redstarts in two distinct habitats, one occupied predominately by males and one mostly by females. All redstarts in both habitat types exhibited similar concentrations of baseline corticosterone levels in fall (October), whereas in spring (March-April), redstarts in female-biased habitat exhibited significantly higher baseline levels regardless of age or sex. In fall, all individuals in both habitats exhibited significant increases in corticosterone concentration with capture and handling, but in spring only redstarts (both sexes) in male-biased habitat continued to exhibit acute corticosterone secretion. Redstarts in female-biased habitat had elevated baseline corticosterone levels and reduced acute corticosterone secretion. In spring, baseline corticosterone concentration was negatively correlated with body mass, suggesting muscle catabolism associated with high corticosterone concentrations or possibly that birds are leaner as a result of increased foraging effort. These results indicate that redstarts (primarily females) in female-biased habitats suffered a decline in physiological condition, which could in turn influence their departure schedules, migration patterns and even their condition and arrival schedules on the breeding grounds. Thus, segregation of populations into habitats of different quality during the non-breeding period may have ramifications throughout the annual cycle of such migratory species. Furthermore, these results show the usefulness of plasma corticosterone levels as indicators of physiological condition and thus habitat quality for birds during the non-breeding period.