The effects of chronic psychological and physical stress on feather replacement in European starlings (Sturnus vulgaris)

Corticosterone (CORT) is seasonally modulated in many passerines, with plasma CORT concentrations lowest during the prebasic molt, when all feathers are replaced. Recent evidence indicating that CORT implants slow the rate of feather regrowth in molting birds suggests that plasma CORT concentrations are downregulated during molt in order to avoid the inhibition of feather growth caused by the protein catabolic activity of CORT. To further test this hypothesis, we examined whether endogenous CORT release, stimulated by exposure to either psychological stress or physical stress (food restriction), could inhibit feather regrowth rates or decrease feather quality in birds undergoing an induced molt (feather replacement after plucking). European starlings (Sturnus vulgaris) were exposed to chronic psychological stress or food restriction for three weeks of the feather regrowth period. Throughout this time, the length of growing primary, secondary, and tail feathers was measured and blood samples were collected to measure baseline and stress-induced CORT concentrations. Upon completion of growth, feather quality was analyzed via measurements of mass, rachis length, feather area, and presence of fault bars. Both psychological and physical stress protocols elevated circulating plasma CORT but significantly less than implants from an earlier study did. Psychological stress had no effect on feather regrowth rates or feather quality. Food restriction had no effect on feather growth rate but caused asynchronous feather replacement. When combined with psychological stress, physical stress also resulted in smaller feather area. Results indicate that CORT implants may not accurately represent chronic stress physiology. Additionally, the purpose for downregulating CORT concentrations during molt appears to be more complicated than simply protecting feather production from CORT's catabolic effects.     

The effects of chronic psychological and physical stress on feather replacement in European starlings (Sturnus vulgaris).

Corticosterone (CORT) is seasonally modulated in many passerines, with plasma CORT concentrations lowest during the prebasic molt, when all feathers are replaced. Recent evidence indicating that CORT implants slow the rate of feather regrowth in molting birds suggests that plasma CORT concentrations are downregulated during molt in order to avoid the inhibition of feather growth caused by the protein catabolic activity of CORT. To further test this hypothesis, we examined whether endogenous CORT release, stimulated by exposure to either psychological stress or physical stress (food restriction), could inhibit feather regrowth rates or decrease feather quality in birds undergoing an induced molt (feather replacement after plucking). European starlings (Sturnus vulgaris) were exposed to chronic psychological stress or food restriction for three weeks of the feather regrowth period. Throughout this time, the length of growing primary, secondary, and tail feathers was measured and blood samples were collected to measure baseline and stress-induced CORT concentrations. Upon completion of growth, feather quality was analyzed via measurements of mass, rachis length, feather area, and presence of fault bars. Both psychological and physical stress protocols elevated circulating plasma CORT but significantly less than implants from an earlier study did. Psychological stress had no effect on feather regrowth rates or feather quality. Food restriction had no effect on feather growth rate but caused asynchronous feather replacement. When combined with psychological stress, physical stress also resulted in smaller feather area. Results indicate that CORT implants may not accurately represent chronic stress physiology. Additionally, the purpose for downregulating CORT concentrations during molt appears to be more complicated than simply protecting feather production from CORT's catabolic effects.     

Elevated corticosterone in feathers correlates with corticosterone‐induced decreased feather quality: a validation study

The newly described technique of extracting corticosterone (CORT) from bird feathers may serve as a less invasive, more integrated measure of a bird's stress response. Previous work indicated that elevated plasma CORT resulted in poorer quality feathers during molt. We tested the hypothesis that a direct link exists between plasma and feather CORT concentrations. We experimentally increased plasma CORT concentrations using implants and found that the corresponding rise in CORT could be detected in feathers grown during implantation. Furthermore, CORT levels in two feathers grown at the same time from the same bird were very consistent. These results provide evidence that elevated CORT is a causative factor in decreasing feather quality during molt. However, there remain technical details that suggest caution when interpreting data from CORT extracted from feathers. Different portions of a growing feather did not necessarily reflect changes in plasma CORT at the time different parts of the feather were forming, a standard pool of homogenized feathers indicated that sample mass affects measured feather CORT concentration, and different antibodies produced different measured CORT concentrations, leaving in doubt the exact steroid being assayed.     

Does Feather Corticosterone Reflect Individual Quality or External Stress in Arctic-Nesting Migratory Birds?

The effects of environmental perturbations or stressors on individual states can be carried over to subsequent life stages and ultimately affect survival and reproduction. The concentration of corticosterone (CORT) in feathers is an integrated measure of hypothalamic–pituitary–adrenal activity during the molting period, providing information on the total baseline and stress-induced CORT secreted during the period of feather growth. Common eiders and greater snow geese replace all flight feathers once a year during the pre-basic molt, which occurs following breeding. Thus, CORT contained in feathers of pre-breeding individuals sampled in spring reflects the total CORT secreted during the previous molting event, which may provide insight into the magnitude or extent of stress experienced during this time period. We used data from multiple recaptures to disentangle the contribution of individual quality vs. external factors (i.e., breeding investment or environmental conditions) on feather CORT in arctic-nesting waterfowl. Our results revealed no repeatability of feather CORT within individuals of either species. In common eiders, feather CORT was not affected by prior reproductive investment, nor by pre-breeding (spring) body condition prior to the molting period. Individual feather CORT greatly varied according to the year, and August-September temperatures explained most of the annual variation in feather CORT. Understanding mechanisms that affect energetic costs and stress responses during molting will require further studies either using long-term data or experiments. Although our study period encompassed only five years, it nonetheless provides evidence that CORT measured in feathers likely reflects responses to environmental conditions experienced by birds during molt, and could be used as a metric to study carry-over effects.     

Nutritional stress affects corticosterone deposition in feathers of Caspian tern chicks

Stressful environmental conditions affect the adrenocortical function of developing animals, which can have consequences for their fitness. Discovery of the avian stress hormone corticosterone (CORT) in feathers has the potential to broaden the application of endocrine research in ecological and evolutionary studies of wild birds by providing a long‐term measure of CORT secretion. Mechanisms of CORT deposition in feathers are not well known and few studies have related feather CORT to circulating plasma CORT during feather growth. Our objective was to experimentally test the validity of using feather CORT as a measure of CORT secretion in developing birds experiencing nutritional stress. Caspian tern Hydroprogne caspia chicks were fed ad libitum or restricted (35% less than ad libitum) diets for four weeks. We measured CORT in feathers from these chicks to examine the relationship between feather CORT concentrations and nutritional limitation, circulating plasma CORT, and feather development. We found that feather CORT was higher in controls fed ad libitum than in restricted individuals, despite higher levels of plasma CORT in restricted chicks compared to controls. Feather mass and growth rates were strongly and positively related to feather CORT concentrations in both treatments. This is the first experimental study to show that feather CORT concentrations can be lower in response to nutritional stress, even when plasma CORT concentrations are elevated. Our results indicate that CORT deposition in feathers may be confounded when feather mass and growth rates are compromised by nutritional stress. We conclude that feather CORT can be used for assessing nutritional stress in growing birds, but the direction of response depends on how strongly stress affects feather development.     

Haste makes waste but condition matters: molt rate-feather quality trade-off in a sedentary songbird

Background

The trade-off between current and residual reproductive values is central to life history theory, although the possible mechanisms underlying this trade-off are largely unknown. The ‘molt constraint’ hypothesis suggests that molt and plumage functionality are compromised by the preceding breeding event, yet this candidate mechanism remains insufficiently explored.

Methodology/Principal Findings

The seasonal change in photoperiod was manipulated to accelerate the molt rate. This treatment simulates the case of naturally late-breeding birds. House sparrows Passer domesticus experiencing accelerated molt developed shorter flight feathers with more fault bars and body feathers with supposedly lower insulation capacity (i.e. shorter, smaller, with a higher barbule density and fewer plumulaceous barbs). However, the wing, tail and primary feather lengths were shorter in fast-molting birds if they had an inferior body condition, which has been largely overlooked in previous studies. The rachis width of flight feathers was not affected by the treatment, but it was still condition-dependent.

Conclusions/Significance

This study shows that sedentary birds might face evolutionary costs because of the molt rate–feather quality conflict. This is the first study to experimentally demonstrate that (1) molt rate affects several aspects of body feathers as well as flight feathers and (2) the costly effects of rapid molt are condition-specific. We conclude that molt rate and its association with feather quality might be a major mediator of life history trade-offs. Our findings also suggest a novel advantage of early breeding, i.e. the facilitation of slower molt and the condition-dependent regulation of feather growth.     

Exogenous and endogenous corticosterone in feathers

In birds, the steroid hormone corticosterone (CORT) increases in response to real or perceived threats to homeostasis. A long‐term record of CORT exposure is recorded in feathers when the hormone is incorporated into the keratinized tissue, and then preserved when the mature feather is cut off from the blood supply. The opportunity to retrospectively assess the exposure of an individual to stressors by measuring the amount of CORT in a feather has generated excitement amongst avian ecologists. However, this technique is relatively new and requires additional validations. In this study, we performed experiments in wild caught European starlings Sturnus vulgaris to test whether: 1) CORT deposition in the feather depends on time of day and 2) whether an ecologically relevant stressor (unpredictable food access) causes a change in feather CORT. We found that exogenous CORT was incorporated into feathers during the day and the night. However, there was no difference in feather CORT between birds with unpredictable access to food and those with continuous access, indicating that feather CORT might not always detect ecologically relevant stressors.     

A Quantitative Analysis of Flight Feather Replacement in the Moustached Tree Swift Hemiprocne mystacea,a Tropical Aerial Forager

The functional life span of feathers is always much less than the potential life span of birds, so feathers must be renewed regularly. But feather renewal entails important energetic, time and performance costs that must be integrated into the annual cycle. Across species the time required to replace flight feather increases disproportionately with body size, resulting in complex, multiple waves of feather replacement in the primaries of many large birds. We describe the rules of flight feather replacement for Hemiprocne mystacea, a small, 60g tree swift from the New Guinea region. This species breeds and molts in all months of the year, and flight feather molt occurs during breeding in some individuals. H. mystacea is one to be the smallest species for which stepwise replacement of the primaries and secondaries has been documented; yet, primary replacement is extremely slow in this aerial forager, requiring more than 300 days if molt is not interrupted. We used growth bands to show that primaries grow at an average rate of 2.86 mm/d. The 10 primaries are a single molt series, while the 11 secondaries and five rectrices are each broken into two molt series. In large birds stepwise replacement of the primaries serves to increase the rate of primary replacement while minimizing gaps in the wing. But stepwise replacement of the wing quills in H. mystacea proceeds so slowly that it may be a consequence of the ontogeny of stepwise molting, rather than an adaptation, because the average number of growing primaries is probably lower than 1.14 feathers per wing.     

Inexplicable Inefficiency of Avian Molt? Insights from an Opportunistically Breeding Arid-Zone Species,Lichenostomus penicillatus

The majority of bird species studied to date have molt schedules that are not concurrent with other energy demanding life history stages, an outcome assumed to arise from energetic trade-offs. Empirical studies reveal that molt is one of the most energetically demanding and perplexingly inefficient growth processes measured. Furthermore, small birds, which have the highest mass-specific basal metabolic rates (BMR_m), have the highest costs of molt per gram of feathers produced. However, many small passerines, including white-plumed honeyeaters (WPHE; Lichenostomus penicillatus), breed in response to resource availability at any time of year, and do so without interrupting their annual molt. We examined the energetic cost of molt in WPHE by quantifying weekly changes in minimum resting metabolic rate (RMR_min) during a natural-molt period in 7 wild-caught birds. We also measured the energetic cost of feather replacement in a second group of WPHEs that we forced to replace an additional 25% of their plumage at the start of their natural molt period. Energy expenditure during natural molt revealed an energy conversion efficiency of just 6.9% (±0.57) close to values reported for similar-sized birds from more predictable north-temperate environments. Maximum increases in RMR_min during the molt of WPHE, at 82% (±5.59) above individual pre-molt levels, were some of the highest yet reported. Yet RMR_min maxima during molt were not coincident with the peak period of feather replacement in naturally molting or plucked birds. Given the tight relationship between molt efficiency and mass-specific metabolic rate in all species studied to date, regardless of life-history pattern (Efficiency (%)  = 35.720•10^−0.494BMRm; r² = 0.944; p = <0.0001), there appears to be concomitant physiological costs entrained in the molt period that is not directly due to feather replacement. Despite these high total expenditures, the protracted molt period of WPHE significantly reduces these added costs on a daily basis.     

Impact of feather molt on ectoparasites: looks can be deceiving

Animals possess a variety of well-documented defenses against ectoparasites, including morphological, behavioral, and immune responses. Another possible defense that has received relatively little attention is the shedding of the host's exterior. The conventional wisdom is that ectoparasite abundance is reduced when birds molt their feathers, mammals molt their hair, and reptiles shed their skin. We carried out an experimental test of this hypothesis for birds by manipulating molt in feral pigeons (Columba livia) infested with feather lice (Phthiraptera: Ischnocera). We used two standard methods, visual examination and body washing, to quantify the abundance of lice on the birds. The visual data indicated a significant effect of molt on lice. However, the more robust body washing method showed that molt had no effect on louse abundance. Two factors caused visual examination to underestimate the number of lice on molting birds. First, molt replaces worn feathers with new, lush plumage that obscures lice during visual examination. Second, we discovered that lice actively seek refuge inside the sheath that encases developing feathers, where the lice cannot be seen. The apparent reduction in louse abundance caused by these factors may account for the conventional wisdom that feather molt reduces ectoparasite abundance in birds. In light of our experimental results, we argue that it is necessary to reinterpret the conclusions of previous studies that were based on observational data. Additional experiments are needed to test whether shedding of the host's exterior reduces ectoparasites in other birds, mammals, and reptiles, similar to the impact of facultative leaf drop on herbivorous insects on trees.

     

An evaluation of feather corticosterone as a biomarker of fitness and an ecologically relevant stressor during breeding in the wild

Feather corticosterone (CORT) levels are increasingly employed as biomarkers of environmental stress. However, it is unclear if feather CORT levels reflect stress and/or workload in the wild. We investigated whether feather CORT represents a biomarker of environmental stress and reproductive effort in tree swallows (Tachycineta bicolor). Specifically, we examined whether individual state and investment during reproduction could predict feather CORT levels in subsequently moulted feathers and whether those levels could predict future survival and reproductive success. Through a manipulation of flight cost during breeding, we also investigated whether an increase in stress level would be reflected in subsequently grown feathers, and whether those levels could predict future success. We found that CORT levels of feathers grown during moult did not (1) reflect past breeding experience (n = 29), (2) predict reproductive output (n = 18), or (3) respond to a manipulation of flight effort during reproduction (10 experimental, 14 control females). While higher feather CORT levels predicted higher return rate (a proxy for survival), they did so only in the manipulated group (n = 36), and this relationship was opposite to expected. Overall, our results add to the mixed literature reporting that feather CORT levels can be positively, negatively, or not related to proxies of within-season and longer-term fitness (i.e., carryover effects). In addition, our results indicate that CORT levels or disturbances experienced during one time (e.g., breeding) may not carry over to subsequent stages (e.g., moult). We, therefore, petition for directed research investigating whether feather CORT represents exposure to chronic stress in feathers grown during moult.     

House sparrows offset the physiological trade‐off between immune response and feather growth by adjusting foraging behavior

Growing feathers and mounting immune responses are both energetically costly for birds. According to the life history trade‐off hypothesis, it has been posited that the costs of feather growth lead to temporal isolation between molt and other expensive activities, reproduction for example. In contrast to life cycle events, the need to mount an immune response can occur at any time, including during feather growth. Thus, we hypothesized that mounting an immune response during feather growth may divert energy and resources from feather growth and impair feather renewal. To test this hypothesis, we clipped or plucked the same feathers of male house sparrows Passer domesticus biblicus. In the clipped group (n = 16), the feathers were absent with no regrowth; in the plucked group (n = 14), feathers were absent and regrowth was initiated. We also had an intact control group of 15 sparrows. We then initiated an inflammatory immune response by subcutaneous injection over the left breast muscle of the birds with a lipopolysaccharide (LPS) and quantified behavioral and physiological responses. We predicted that sparrows with plucked feathers would incur the highest energetic costs while mounting an immune response, and would increase their foraging effort to offset this cost. We found no difference in body mass and resting metabolic rates among sparrows subjected to the different feather and immune treatments. However, we did find that while sparrows with plucked feathers increased foraging efficiency following the immune challenge by paying fewer but longer visits to the food tray, allowing them to maintain food consumption. Foraging efficiency in sparrows with clipped feathers was reduced. We also found that quality of newly grown feathers after the immune challenge was poorer in the plucked group, suggesting that mounting an immune response competes with feather growth for resources.     

Malaria infection negatively affects feather growth rate in the house sparrow Passer domesticus

Birds often face various stressors during feather renewing, for example, enduring infection with blood parasites. Because nutritional resources are typically limited, especially for wild animals, when an individual allocates energy to one physiological system, there is subsequently less for other processes, thereby requiring a trade‐off. Surprisingly, potential trade‐offs between malaria infection and feather growth rate have not been experimentally considered yet. Here, we conducted three studies to investigate whether a trade‐off occurs among feather growth rate, malaria infection and host health conditions. First, we explored whether naturally infected and uninfected house sparrows differed in feather growth rate in the wild. Second, we asked whether experimental inoculation of malaria parasites and/or forcing the renewal of a tail feather. Lastly, we evaluated whether individual condition was affected by experimentally‐induced feather regrowth and/or malaria experimental infection. Our findings showed that feather growth rate was negatively affected by natural malaria infection status in free‐living birds and by experimental infection in captive birds. Furthermore, birds that did not increase body mass or hematocrit during the experimental study had slower feather growth. Together our results suggest that infection with blood parasites has more negative health effects than the growth of tail feathers and that these two processes (response to blood parasite infection and renewal of feathers) are traded‐off against each other. As such, our results highlight the role of malaria parasites as a potential mechanism driving other trade‐offs in wild passerines.     

The Control of Color in Birds

SYNOPSIS. The colors of birds result from deposition of pigments—mainlymelanins and carotenoids—in integumentary structures,chiefly the feathers. The plumages of birds indicate their age,sex, and mode of living, and play important roles in camouflage,mating, and establishment of territories. Since feathers aredead structures, change of color of feathers is effected throughdivestment (molt) and replacement. The color and pattern ofa feather are determined by the interplay of genetic and hormonalinfluences prevailing in its base during regeneration. Mostbirds replace their feathers at least once annually. Some wearthe same kind of basic plumage all the time butothers alternatea basic and breeding plumage, either in one (the male) or bothsexes. Still others may have more than two molts, adding supplementalplumage at certain times in the plumage cycle. The varietiesof patterns of molt, the kinds of plumage, and the colors andpatterns of feathers among birds apparently are the result ofseveral kinds of selection pressures working through evolution.     

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla . As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 98–105.     

The phenology of molting,breeding and their overlap in central Amazonian birds

The energetically challenging periods of molting and breeding are usually temporally separated in temperate birds, but can occur simultaneously in tropical birds, a condition known as molt–breeding overlap. Here, we document great variation in the timing and duration of molting and breeding, and in the extent of molt–breeding overlap, among 87 species of understory passerines in central Amazonia. We analyzed molt and breeding from 26 871 birds captured over a 30‐yr period near Manaus, Brazil. Although most species typically bred during the late dry season (about October through January), many thamnophilids apparently bred year‐round, whereas a few other species from a variety of families bred mainly during the wet season (about January through May). Of all breeding birds with an active brood patch, 12.7% were simultaneously molting. Molt–breeding overlap was more frequently observed among suboscines (13.3%), especially thamnophilids (23.0%), than oscines (6.4%). Some families had <5% molt–breeding overlap frequency, including Tyrannidae (4.4%), Tityridae (0.0%), Pipridae (1.5%), Turdidae (0.0%), and Thraupidae (0.0%), indicating that not all tropical species exhibit molt–breeding overlap. Among 31 well‐sampled species (n ≥15 brood patches), variation in molt–breeding overlap frequency was positively correlated with each species’ average duration of flight feather replacement (range 98–301 d). We also measured feather growth rates of individual birds in nine species; in five of these, slower‐growing feathers increased with an individual's probability of having molt–breeding overlap. Among furnariids, molt–breeding overlap occurred either at the beginning or end of the molt cycle, suggesting that physiological mechanisms typically separate molting from breeding. Thamnophilids showed a much different pattern; molt–breeding overlap occurred at any stage of feather replacement, apparently not regulated to be independent of breeding. These results reveal substantial life‐history variation among Amazonian birds. Future work to resolve the physiological regulation of molting and breeding in tropical birds will greatly contribute to understanding these patterns and their relevance to avian diversity.     

A Trade-Off between Reproduction and Feather Growth in the Barn Swallow (Hirundo rustica)

Physiological trade-offs mediated by limiting energy, resources or time constrain the simultaneous expression of major functions and can lead to the evolution of temporal separation between demanding activities. In birds, plumage renewal is a demanding activity, which accomplishes fundamental functions, such as allowing thermal insulation, aerodynamics and socio-sexual signaling. Feather renewal is a very expensive and disabling process, and molt is often partitioned from breeding and migration. However, trade-offs between feather renewal and breeding have been only sparsely studied. In barn swallows (Hirundo rustica) breeding in Italy and undergoing molt during wintering in sub-Saharan Africa, we studied this trade-off by removing a tail feather from a large sample of individuals and analyzing growth bar width, reflecting feather growth rate, and length of the growing replacement feather in relation to the stage in the breeding cycle at removal and clutch size. Growth bar width of females and length of the growing replacement feather of both sexes were smaller when the original feather had been removed after clutch initiation. Importantly, in females both growth bar width and replacement feather length were negatively predicted by clutch size, and more strongly so for large clutches and when feather removal occurred immediately after clutch completion. Hence, we found strong, coherent evidence for a trade-off between reproduction, and laying effort in particular, and the ability to generate new feathers. These results support the hypothesis that the derived condition of molting during wintering in long-distance migrants is maintained by the costs of overlapping breeding and molt.     

Prebasic molt initiation and progress in northern fulmars of the High Arctic: do molt and breeding overlap?

We examined feather molt progress of northern fulmars (Fulmarus glacialis) at Cape Vera in the Canadian High Arctic through opportunistic observation of individuals in flight from 2003 to 2006, and examination of bodies and wings of 127 individuals collected at the site, from 2003 to 2005. We found no evidence suggesting that fulmars shed primary feathers during breeding. Prebasic molt was initiated in the head, neck, sides, belly and back approximately 1 week before hatch. We failed to detect a sex effect on molt progress, but molt among breeders was delayed compared to molt in non- or failed breeders. This study constitutes a baseline we feel may be useful to: (1) researchers interested in feather replacement chronology, wherein feathers are used as sources of biological information; and (2) researchers interested in eventual assessment of relationships among large-scale environmental processes and molt progress in this species, especially in light of predicted changes to Arctic regions.     

Feather isotope analysis discriminates age-classes of Western, Least, and Semipalmated sandpipers when plumage methods are unreliable

ABSTRACT Avian age‐class discrimination is typically based on the completeness of the first prebasic molt. In several calidrid sandpiper species, juvenal flight feathers grown on Arctic breeding grounds are retained through the first three migrations. Thereafter, flight feathers are grown annually at temperate migratory stopover sites during the fall or on the subtropical wintering grounds. Standard methods for distinguishing age classes of sandpipers rely on a combination of traits, including body plumage, coloration of protected inner median covert edges, and extent of flight feather wear. We tested the ability of stable hydrogen isotope ratios in flight feathers (δD_f) to distinguish young birds in their first winter through second fall from older adults in three calidrid sandpiper species, Western (Calidris mauri), Least (C. minutilla), and Semipalmated (C. pusilla) sandpipers. We compared the apparent reliability of the isotope approach to that of plumage‐based aging. The large expected differences in δD_f values of flight feathers grown at Arctic versus non‐Arctic latitudes enabled use of this technique to discriminate between age‐classes. We determined δD_f values of known Arctic‐grown feathers from juveniles that grew their flight feathers on the breeding grounds. Flight feather δD_f values of southward‐migrating adults showed bimodal distributions for all three species. Negative values overlapped with species‐specific juvenile values, identifying putative second fall birds with high‐latitude grown juvenal feathers retained from the previous year. The more positive values identified older adults who grew their feathers at mid‐ and low latitudes. Importantly, δD_f analysis successfully identified first‐winter and second‐fall birds not detected by plumage‐based aging. Flight feather wear alone was a poor basis for age classification because scores overlapped extensively between putative second fall birds and older adults. Flight feather hydrogen isotope analysis enables more definitive assignment of age classes when standard plumage methods are unreliable.     

Continent-wide variation in feather colour of a migratory songbird in relation to body condition and moulting locality

Understanding the causes of variation in feather colour in free-living migratory birds has been challenging owing to our inability to track individuals during the moulting period when colours are acquired. Using stable-hydrogen isotopes to estimate moulting locality, we show that the carotenoid-based yellow-orange colour of American redstart (Setophaga ruticilla) tail feathers sampled on the wintering grounds in Central America and the Caribbean is related to the location where feathers were grown the previous season across North America. Males that moulted at southerly latitudes were more likely to grow yellowish feathers compared with males that moulted more orange-red feathers further north. Independent samples obtained on both the breeding and the wintering grounds showed that red chroma-an index of carotenoid content-was not related to the mean daily feather growth rate, suggesting that condition during moult did not influence feather colour. Thus, our results support the hypothesis that feather colour is influenced by ecological conditions at the locations where the birds moulted. We suggest that these colour signals may be influenced by geographical variation in diet related to the availability of carotenoids.