Feather corticosterone is lower in translocated and historical populations of the endangered Laysan duck (Anas laysanensis).

Identifying reliable bioindicators of population status is a central goal of conservation physiology. Physiological stress measures are often used as metrics of individual health and can assist in managing endangered species if linked to fitness traits. We analysed feather corticosterone, a cumulative physiological stress metric, of individuals from historical, translocated, and source populations of an endangered endemic Hawaiian bird, the Laysan duck (Anas laysanensis). We hypothesized that feather corticosterone would reflect the improved reproduction and survival rates observed in populations translocated to Midway and Kure Atolls from Laysan Island. We also predicted less physiological stress in historical Laysan birds collected before ecological conditions deteriorated and the population bottleneck. All hypotheses were supported: we found lower feather corticosterone in the translocated populations and historical samples than in those from recent Laysan samples. This suggests that current Laysan birds are experiencing greater physiological stress than historical Laysan and recently translocated birds. Our initial analysis suggests that feather corticosterone may be an indicator of population status and could be used as a non-invasive physiological monitoring tool for this species with further validation. Furthermore, these preliminary results, combined with published demographic data, suggest that current Laysan conditions may not be optimal for this species.

Research Note: Irritating flashing light or poultry-friendly lighting-are flicker frequencies of LED luminaires a potential stress factor in the husbandry of male fattening turkeys?

Conventional fluorescent tubes are increasingly being replaced with innovative light-emitting diodes (LEDs) for lighting poultry houses. However, little is known about whether the flicker frequencies of LED luminaires are potential stressors in poultry husbandry. The term "light flicker" describes the fluctuations in the brightness of an electrically operated light source caused by the design and/or control of the light source. In this context, the critical flicker frequency (CFF) characterizes the frequency at which a sequence of light flashes is perceived as continuous light. It is known that CFF in birds is higher than that in humans and that light flicker can affect behavioral patterns and stress levels in several bird species. As there is a lack of knowledge about the impact of flicker frequency on fattening turkeys, this study aimed to investigate the effects of flicker frequency on the behavior, performance, and stress response in male turkeys. In 3 trials, a total of 1,646 male day-old turkey poults of the strain B.U.T. 6 with intact beaks were reared for 20 wk in 12 barn compartments of 18 m² each. Each barn compartment was illuminated using 2 full-spectrum LED lamps. Flicker frequencies of 165 Hz, 500 Hz, and 16 kHz were set in the luminaires to illuminate the compartments. Analyses of feather corticosterone concentration were performed on fully grown third-generation primaries (P 3) of 5 turkeys from each compartment. No significant differences were found in the development of live weight, feed consumption, or prevalence of injured or killed turkeys by conspecifics reared under the above flicker frequencies. The flicker frequencies also did not significantly influence feather corticosterone concentrations in the primaries of the turkeys. In conclusion, the present results indicate that flicker frequencies of 165 Hz or higher have no detrimental effect on growth performance, injurious pecking, or endocrine stress response in male turkeys and, thus, may be suitable for use as animal-friendly lighting.

Long term stability of corticosterone in feathers.

Measuring corticosterone in feathers allows researchers to make long-term, retrospective assessments of physiology with non-invasive sampling. To date, there is little evidence that steroids degrade within the feather matrix, however this has yet to be determined from the same sample over many years. In 2009, we made a pool of European starling (Sturnus vulgaris) feathers that had been ground to a homogenous powder using a ball mill and stored on a laboratory bench. Over the past 14 years, a subset of this pooled sample has been assayed via radioimmunoassay (RIA) 19 times to quantify corticosterone. Despite high variability across time (though low variability within assays), there was no effect of time on measured feather corticosterone concentration. In contrast, two enzyme immunoassays (EIA) produced higher concentrations than the samples assayed with RIA, though this difference is likely due to different binding affinities of the antibodies used. The present study provides further support for researchers to use specimens stored long-term and from museums for feather corticosterone quantification, and likely applies to corticosteroid measurements in other keratinized tissues.

Effects of Housing System on Anxiety, Chronic Stress, Fear, and Immune Function in Bovan Brown Laying Hens.

The scientific community needs objective measures to appropriately assess animal welfare. The study objective was to assess the impact of housing system on novel physiological and behavioral measurements of animal welfare for laying hens, including secretory and plasma Immunoglobulin (IgA; immune function), feather corticosterone (chronic stress), and attention bias testing (ABT; anxiety), in addition to the well-validated tonic immobility test (TI; fearfulness). To test this, 184 Bovan brown hens were housed in 28 conventional cages (3 birds/cage) and 4 enriched pens (25 birds/pen). Feces, blood, and feathers were collected 4 times between week 22 and 43 to quantify secretory and plasma IgA and feather corticosterone concentrations. TI tests and ABT were performed once. Hens that were from cages tended to show longer TI, had increased feather corticosterone, and decreased secretory IgA at 22 weeks of age. The caged hens fed quicker, and more hens fed during the ABT compared to the penned hens. Hens that were in conventional cages showed somewhat poorer welfare outcomes than the hens in enriched pens, as indicated by increased chronic stress, decreased immune function at 22 weeks of age but no other ages, somewhat increased fear, but reduced anxiety. Overall, these novel markers show some appropriate contrast between housing treatments and may be useful in an animal welfare assessment context for laying hens. More research is needed to confirm these findings.

Hungry or angry? Experimental evidence for the effects of food availability on two measures of stress in developing wild raptor nestlings.

Food shortage challenges the development of nestlings; yet, to cope with this stressor, nestlings can induce stress responses to adjust metabolism or behaviour. Food shortage also enhances the antagonism between siblings, but it remains unclear whether the stress response induced by food shortage operates via the individual nutritional state or via the social environment experienced. In addition, the understanding of these processes is hindered by the fact that effects of food availability often co-vary with other environmental factors. We used a food supplementation experiment to test the effect of food availability on two complementary stress measures, feather corticosterone (CORTf) and heterophil/lymphocyte ratio (H/L) in developing red kite (Milvus milvus) nestlings, a species with competitive brood hierarchy. By statistically controlling for the effect of food supplementation on the nestlings' body condition, we disentangled the effects of food and ambient temperature on nestlings during development. Experimental food supplementation increased body condition, and both CORTf and H/L were reduced in nestlings of high body condition. Additionally, CORTf decreased with age in non-supplemented nestlings. H/L decreased with age in all nestlings and was lower in supplemented last-hatched nestlings compared with non-supplemented ones. Ambient temperature showed a negative effect on H/L. Our results indicate that food shortage increases the nestlings' stress levels through a reduced food intake affecting both their nutritional state and their social environment. Thus, food availability in conjunction with ambient temperature shapes between- and within-nest differences in stress load, which may have carry-over effects on behaviour and performance in further life-history stages.

Cooling increases corticosterone deposition in feathers of eastern bluebird chicks.

Exposure to noxious stimuli early in life can both activate and shape the development of the hypothalamic-pituitary-adrenal (HPA) axis in birds and other vertebrates, with the potential for lifelong consequences. Studies assessing early HPA axis activation often rely on collection of blood samples to evaluate circulating glucocorticoid levels. However, blood sampling in small altricial young is invasive, limited by animal size, and not sufficient to provide detailed information about hormone exposure over protracted periods of time. We tested the use of feather corticosterone as an alternative method to assess HPA axis activity early in life in free-living, altricial chicks, for whom timing of growth of first feathers coincides with a period of rapid growth, development of the HPA axis, and reliance on parental care. We investigated (1) whether ecologically relevant bouts of experimental cooling prior to the onset of homeothermy-conditions known to elevate circulating corticosterone-are reflected in changes of feather corticosterone deposition in Eastern bluebird (Sialia sialis) chicks, and (2) whether such changes occurred in a sex-dependent manner. We found that cooling during the first week of life resulted in elevated feather corticosterone in first-grown feathers of experimentally cooled chicks relative to controls. The timing of deposition of corticosterone in feathers in response to temperature treatments was delayed in females compared to males. Results indicate that the hormone deposition in feather tissues of altricial nestlings reflects exposure to environmental stimuli, and can thus provide a minimally invasive tool for assessing HPA activity in early life. The development of the HPA axis, or its activation in response to environmental stimuli early in life, may also occur in a sex-dependent manner in altricial birds.

Variation in feather corticosterone levels in Alpine swift nestlings provides support for the hypo-responsive hypothesis.

In birds, feather corticosterone values (CORTf) are increasingly used as a retrospective and integrative proxy of an individual's physiological state during the period of feather growth. Relatively high CORTf values are usually interpreted as an indicator of exposure to energy-demanding or stressful conditions during feather growth. However, in nestlings this interpretation might not always hold true. The reasons are that, firstly nestlings (especially altricial ones) still develop their hypothalamo-pituitary-adrenal (HPA) reactivity during the growth of their feathers. Hence, at a young age, nestlings might be unable to mount a substantial adrenocortical stress response. Secondly, some species are able to down-regulate their metabolism during food scarcity and therewith probably also their CORT release. Consequently, CORTf values may not unambiguously reflect whether nestlings have suffered from energy-demanding or stress situations. Relatively high CORTf values might indicate either energy-demanding or stressful conditions ('stress responsive hypothesis'), or - conversely - favourable conditions during the period of feather growth ('hypo-responsive hypothesis'). In the altricial Alpine swift (Tachymarptis melba), we tested which factors help to distinguish between the two hypotheses by considering factors which affect CORT release (brood size, weather) and factors which are affected by high CORT levels (nestling size and condition). We measured CORTf in 205 nestlings over 7 years and collected data on brood size, body size, body condition and prevailing weather. Nestling CORTf values were positively correlated with body condition and negatively with adverse weather, supporting the hypo-responsive hypothesis. Results from the Alpine swift study, supplemented with a survey of the literature, show that relatively easily collected parameters on brood size, nestling size and condition, and environmental factors can help to distinguish between the two hypotheses. A meaningful interpretation of nestling CORTf should only be made in the context of species-specific traits.

Interannual measures of nutritional stress during a marine heatwave (the Blob) differ between two North Pacific seabird species.

'The Blob', a mass of anomalously warm water in the Northeast Pacific Ocean peaking from 2014 to 2016, caused a decrease in primary productivity with cascading effects on the marine ecosystem. Among the more obvious manifestations of the event were seabird breeding failures and mass mortality events. Here, we used corticosterone in breast feathers (fCort), grown in the winter period during migration, as an indicator of nutritional stress to investigate the impact of the Blob on two sentinel Pacific auk species (family Alcidae). Feathers were collected from breeding females over 8 years from 2010 to 2017, encompassing the Blob period. Since Pacific auks replace body feathers at sea during the migratory period, measures of fCort provide an accumulated measure of nutritional stress or allostatic load during this time. Changes in diet were also measured using δ15N and δ13C values from feathers. Relative to years prior to the Blob, the primarily zooplanktivorous Cassin's auklets (Ptychoramphus aleuticus) had elevated fCort in 2014-2017, which correlated with the occurrence of the Blob and a recovery period afterwards, with relatively stable feather isotope values. In contrast, generalist rhinoceros auklets (Cerorhinca monocerata) displayed stable fCort values across years and increased δ15N values during the Blob. As marine heatwaves increase in intensity and frequency due to climate change, this study provides insight into the variable response of Pacific auks to such phenomena and suggests a means for monitoring population-level responses to climatological variation.

Comparison of Two Different Feather Sampling Methods to Measure Corticosterone in Wild Greater Flamingos (Phoenicopterus roseus) and Wild Mallards (Anas platyrhynchos).

This research project had the aim to validate the possible alternative and less-painful sampling method of cutting feathers close to the skin instead of plucking them for subsequent feather corticosterone analysis, confirming recently-published results for other species in captivity. Analyzing CORTf is often used in animal welfare studies in combination with behavioral monitoring. The background of this idea was to act in the sense of animal welfare and reduce the burden of animal studies according to the 3-R-Principle (Replacement, Reduction, and Refinement) by refining procedures. To confirm the hypothesis that the sampling method itself has no influence on CORTf levels measured, plucked and cut samples of the respective bird were collected. Birds of two wild species were used: the Mallard (Anas platyrhynchos) and the Greater Flamingo (Phoenicopterus roseus). The CORTf was measured by using an enzyme-linked immunosorbent assay (ELISA). The determined values were inspected for their mean values, standard deviation (SD), and average differences. Afterwards, the CORTf levels of both species were compared, according to the sampling method, with the concordance correlation coefficient (CCC). In the Bland-Altman (BA) plot the differences of the methods were displayed against the mean values. Additionally, sex, as a possible factor influencing CORTf, was analyzed using the Mann-Whitney U test. The values of CCC showed poor agreement in the comparability of the two methods, whereas the concordance of the BA plot was decent. The average differences between the methods were marginal for both species (Mallards: -0.16 pg/mm, Flamingos -0.13 pg/mm). In summary, all anomalies or differences between the methods were negligible. Therefore, the alternative sampling method seems to be as suitable as the common standard method. No significant difference was found between females and males. Nevertheless, our results suggest that CORTf should not be interpreted in just considering the values themselves, but the results they should be analyzed in the context of a wider set of parameters. Hence, further studies are encouraged to create a larger data pool.

Feather Corticosterone Measurements and Behavioral Observations in the Great White Pelican (Pelecanus onocrotalus) Living under Different Flight Restraint Conditions in German Zoos.

The pinioning of birds was previously one of the most-accepted forms of mutilation in zoos. Despite a lack of knowledge on the effects of deflighting procedures with regard to the well-being of deflighted birds, pelicans are often reversibly deflighted by feather-clipping to keep them in open enclosures, including those with ponds without netting. In the present study, we focused on the welfare implications of flight restraint on one of the most commonly kept types of birds in German zoos, the great white pelican. A combination of behavioral observations and feather corticosterone concentrations (CORTf) of pelicans with different deflighting statuses (i.e., irreversibly deflighted, reversibly deflighted, and airworthy) was used to evaluate the effects of deflighting status on pelican welfare. We observed 215 individuals in 21 different German zoos. The pelicans lived in differently designed exhibits. An ethogram for these species was developed and their behavior was evaluated by scan sampling. Feather samples from 182 individuals were collected to determine if different deflighting conditions influenced the CORTf and therefore stress levels. The hypothesis was that the CORTf values of airworthy pelicans differ from those of deflighted pelicans. Tendencies with regard to the flight status groups were found. Conversely, reversibly deflighted pelicans had higher CORTf levels than irreversible deflighted and airworthy pelicans. Tendencies with regard to CORTf values and the group size of the kept pelicans were observed. The CORTf values were lower in groups consisting of more than 10 animals. In addition, the frequency of fluttering behavior was positively associated with CORTf values. Pelicans that frequently showed fluttering had higher CORTf values. Therefore, fluttering behavior might be considered a sign of stress levels in pelicans. This study is one of the first important steps in assessing the impact of deflighting procedures on the welfare of great white pelicans kept in zoos.

Research Note: A sip of stress. Effects of corticosterone supplementation in drinking water on feather corticosterone concentrations in layer pullets.

The measurement of feather corticosterone concentrations (fCORT) is a comparatively new method for the evaluation of stress in wild and captive birds and may be a useful indicator in animal welfare research. The aim of this study was to assess the suitability of fCORT as an indicator of stress, and for this purpose a prolonged stress period was experimentally simulated by oral intake of corticosterone via drinking water and corticosterone concentrations were analyzed in feathers grown during this period. Layer pullets of both a control group (n = 20) and a CORT group (n = 20) were offered drinking water ad libitum throughout the entire experimental phase. The drinking water of the CORT group was supplemented with corticosterone at a concentration of 20 mg/l from the 64th to the 114th day of life. The vaned parts of the primaries 5 (P5s) were clipped on d 114 and fCORT was analyzed by ELISA after extraction. Body weights increased from day 64 until d 114 in both groups, however, at the end of the experiment, mean body mass in the CORT group was significantly lower than in the control group (P < 0.001). Pullets of the CORT group also showed shorter and lighter P5s as well as a retarded molt of the primaries. The supplementation of drinking water with corticosterone increased the average fCORT in the P5s of the CORT pullets compared with the control group (median: 110.3 pg/mm [interquartile range (IQR): 47.2] vs. 10.0 pg/mm [IQR: 2.5], P < 0.001). The results show that experimentally increased systemic corticosterone concentrations over a period of seven weeks in layer pullets are reflected in corticosterone concentrations of feathers grown during that time. This indicates that the measurement of fCORT may be a useful and minimally invasive tool for the evaluation of long-term stress in chicken and provides the basis for further investigations on its use in animal welfare research.

Selection on individuals of introduced species starts before the actual introduction.

Biological invasion is a global problem with large negative impacts on ecosystems and human societies. When a species is introduced, individuals will first have to pass through the invasion stages of uptake and transport, before actual introduction in a non-native range. Selection is predicted to act during these earliest stages of biological invasion, potentially influencing the invasiveness and/or impact of introduced populations. Despite this potential impact of pre-introduction selection, empirical tests are virtually lacking. To test the hypothesis of pre-introduction selection, we followed the fate of individuals during capture, initial acclimation, and captivity in two bird species with several invasive populations originating from the international trade in wild-caught pets (the weavers Ploceus melanocephalus and Euplectes afer). We confirm that pre-introduction selection acts on a wide range of physiological, morphological, behavioral, and demographic traits (incl. sex, age, size of body/brain/bill, bill shape, body mass, corticosterone levels, and escape behavior); these are all traits which likely affect invasion success. Our study thus comprehensively demonstrates the existence of hitherto ignored selection acting before the actual introduction into non-native ranges. This could ultimately change the composition and functioning of introduced populations, and therefore warrants greater attention. More knowledge on pre-introduction selection also might provide novel targets for the management of invasive species, if pre-introduction filters can be adjusted to change the quality and/or quantity of individuals passing through such that invasion probability and/or impacts are reduced.

Validation of an Alternative Feather Sampling Method to Measure Corticosterone.

The most common feather sampling method for feather corticosterone measurement is by plucking the feathers from the bird's skin. This procedure performed on living, restrained birds is qualified as an animal experiment according to German/European legislation, which has to be applied for from the competent authorities. The Directive 2010/63/EU requires the full implementation of the 3-R Principle of Russel and Burch in animal experiments, which means not only to replace the use of animals, but also to reduce the number of animals used and to refine procedures whenever possible. In response to this issue, the aim of this study was to validate an alternative, less invasive sampling method by cutting feathers close to the skin in comparison to the gold standard of plucking them. For this proof-of-principle study, a conventional poultry husbandry with trial groups of geese (Anser anser domesticus) and ducks (Anas sterilis) was selected. All birds were kept under the same living conditions to standardize the influencing factors regarding husbandry, and thus, their stress levels. Feather samples were collected between the shoulders from 46 geese and 51 ducks, both by cutting as well as by plucking, directly after slaughter for meat production. Feather corticosterone levels were measured with Enzyme-Linked Immunosorbent Assay (ELISA). Results were compared using Bland-Altman plots and concordance correlation coefficients (CCC). It could be seen that concordance between corticosterone levels in cut and plucked feathers was rather poor: 0.38 for Anser, and 0.57 for Anas. However, comparing the mean corticosterone values in pg/mm of each species with their respective standard deviations, the differences between the methods were negligible. As the results showed that the differences between the individuals were markedly greater than the differences between the methods, the determination of corticosterone levels in cut feathers is valid compared to using plucked feathers. The validation tests of ELISA showed only acceptable repeatability and reliability. Hence, the results should be verified in further studies. In conclusion, it is recommended for future research to use cut instead of plucked feathers for corticosterone measurement.

Research Note: Quantifying corticosterone in turkey (Meleagris gallopavo) feathers using ELISA.

The measurement of corticosterone in feathers is an appealing tool for assessing glucocorticoids in wild and domestic bird species. Feather corticosterone measurements can be performed noninvasively and can provide a means for comparing glucocorticoid secretion between individual birds; thus, such measurements can be used to assess the welfare of domestic poultry. The focus of this study was to assess the validity of detecting corticosterone in turkey (Meleagris gallopavo) feathers using an ELISA and investigate differences between genetic lines. Primary feather 9 was obtained at a processing plant from 123 individuals from 3 different purebred turkey lines (line A [N = 46], line B [N = 24], and line C [N = 53]). Assay validation tests were performed using a commercially available ELISA kit (Cayman Chemicals, Cedarlane Labs, Canada). Indicators of accuracy, recovery, precision, and sensitivity were sufficient. Significant differences in feather corticosterone concentration between the 3 lines were observed. Line C had significantly higher feather corticosterone than line A (P < 0.0001) and line B (P = 0.036). These results indicate that the quantification of feather corticosterone using an ELISA is a valid method for assessing glucocorticoid levels in turkeys. This is the first report of differences in feather corticosterone between different purebred turkey lines. Differences observed between purebred lines provide an intriguing basis for further investigation into the genetic parameters of glucocorticoid levels in turkeys.

Feather Corticosterone Measurements of Greater Flamingos Living under Different Forms of Flight Restraint.

Deflighting zoo birds is a practice that receives increasing criticism due to its presumed incompatibility with animal welfare. To our knowledge, this is the first approach to address this problem in a scientific way. To do this, we compared feather corticosterone (CORTf) from Greater Flamingos (Phoenicopterus roseus, n = 151) of different flight status (i.e., pinioned, feather clipped, airworthy) from twelve different zoological institutions. To complement the hormone measurements, behavioral observations (scan samplings) were conducted prior to feather sampling within the presumed time frame of feather growth. We hypothesized that CORTf of the deflighted flamingos would differ from CORTf of their airworthy conspecifics. No significant difference in CORTf was found between the three groups, and our hypothesis was rejected. However, the impact of the institution itself (i.e., the housing conditions) proved to be the most dominant variable (variance between the institutions = 53.82%). Due to high variability, the behavioral observations were evaluated descriptively but did not give rise to doubt the findings in CORTf. Therefore, we assume that the method of flight restraint of Greater Flamingos does not have a measurable effect on CORTf. We consider this model for evaluating animal welfare of zoo birds a useful tool and provide ideas for further adjustments for consecutive studies.

Effects of Dark Brooder Rearing and Age on Hypothalamic Vasotocin and Feather Corticosterone Levels in Laying Hens.

Chickens cannot independently thermoregulate at hatch and lack opportunity to behaviorally thermoregulate with a hen in the egg layer industry, thus barns are heated to thermoneutral temperatures. Dark brooders are low-energy-consuming hot plates, which may be environmentally advantageous while providing welfare-enhancing aspects of maternal care (i.e., shelter and separation of active and inactive individuals). Dark brooder use has been demonstrated to decrease injurious pecking and mortality well into the production period of layers. To further understand hen development around lay onset and effects of dark brooders on the brain and HPA-axis, we examined effects of rearing with dark brooders on expression of vasotocin (AVT) in the hypothalamus and corticosterone (CORT) in the feathers of in total 48 layer Isa Warren hens at 16 w and 28 w of age (n = 12 per age and treatment). An age-dependent decreased number of AVT-positive neurons was seen in the medial preoptic area, medial preoptic nucleus, paraventricular nucleus, rostral part (prepeduncular hypothalamus), and lateral preoptic area. Trends to effects of brooder rearing were found in both anteromedial preoptic nucleus and supraoptic nucleus, with dark brooder reared animals showing higher mean counts of AVT-positive neurons in both areas. No interactions between brooder raising and age were observed in AVT-positive neuron count. CORT levels were higher in primary wing feathers from 28 week old hens than in those from 16 week hens. No main effects of rearing with dark brooders or interactions between age and treatment were found on CORT levels. The age-dependent effects seen in the hypothalamus and CORT aids in further understanding of the development of chickens around puberty. The use of brooders tended to increase AVT expression in the anteromedial preoptic nucleus and supraoptic nucleus, an indication that dark brooder rearing may affect physiological responses regulated by these areas. The lack of effect of dark brooders on CORT in feathers is at the least an indication that the use of dark brooders is not stressful; in combination with the benefits of dark brooders on injurious pecking, fearfulness and early mortality, this pleads for the use of dark brooders in on-farm situations.

Raising young with limited resources: supplementation improves body condition and advances fledging of Canada Jays.

Food availability early in life can play a vital role in an individual's development and success, but experimental evidence for the direct effects of food on body condition, physiology, and survival of young animals in the wild is still relatively scarce. Food-caching Canada Jays (Perisoreus canadensis) begin breeding in the late winter and, therefore, rely on either cached food or seemingly limited quantities of fresh food to feed nestlings in the early spring. Using a 2-yr food supplementation experiment conducted during the nestling period and 40 yr of observational data on food supplemented by the public, we examined whether food quantity during early life influenced the physiology, body condition, timing of fledging, and survival of young Canada Jays in Algonquin Provincial Park, Ontario, Canada. Experimental food supplementation of breeding pairs advanced the fledging date of young by 24% (5.5 d) compared to controls. In 1 yr of the experiment, nestlings raised on experimentally supplemented territories had lower feather corticosterone concentrations and were in higher body condition than controls. Across treatment and control nests, young that successfully fledged had lower concentrations of feather corticosterone and were in higher body condition than those that did not fledge. Based on 40 yr of observational data, nestling body condition was positively related to the degree of food supplementation by park visitors and nestlings in higher body condition were more likely to be observed in the population in the following fall. Our results demonstrate how food availability early in life can have important downstream consequences on metrics related to individual fitness, including first year survival.

Corticosterone levels correlate in wild-grown and lab-grown feathers in greenfinches (Carduelis chloris) and predict behaviour and survival in captivity.

Level of corticosterone (CORT), which is a predominant glucocorticoid in birds, has become the main indicator for assessing the stress level of birds in ecological studies. Feather corticosterone (CORTf) provides information about corticosterone levels during feather growth, however, the underlying causes of individual variation of CORTf between individuals and individual persistency of CORTf are not yet fully understood. Therefore, this study addresses individual consistency in CORTf and the association of variation in CORTf with behaviour that results in damage to tail feathers. We studied relations between CORTf, plasma CORT, and behaviour in wild-caught male greenfinches in captivity. CORTf in wild-grown feathers correlated positively with CORTf in lab-grown feathers. CORTf levels were about 20% lower in lab-grown feathers than in those grown in the wild. Four birds that died in captivity had significantly higher average CORTf levels in wild-grown feathers than the survivors. Plasma CORT levels of two measurements taken in the lab seven days apart correlated positively, however, no correlations between plasma CORT and CORTf were found. In order to study the link between CORTf and behaviour, the extent of tail damage from flapping against cage bar was assessed. Contrary to our prediction, birds with higher CORTf had less tail damage. This study adds to the evidence that CORTf levels can be considered as informative markers of some persistent component of individual phenotypic quality that can predict survival under standardized laboratory conditions.

Sex-specific effects of the in ovo environment on early-life phenotypes in eiders.

Maternal effects affect offspring phenotype and fitness. However, the roles of offspring sex-specific sensitivity to maternal glucocorticoids and sex-biased maternal investment remain unclear. It is also uncertain whether telomere length (a marker associated with lifespan) depends on early growth in a sex-specific manner. We assessed whether maternal traits including corticosterone (CORT; the main avian glucocorticoid) and in ovo growth rate are sex-specifically related to offspring CORT exposure, relative telomere length (RTL) and body condition in eiders (Somateria mollissima). We measured feather CORT (fCORT), RTL and body condition of newly hatched ducklings, and growth rate in ovo was expressed as tarsus length at hatching per incubation duration. Maternal traits included baseline plasma CORT, RTL, body condition and breeding experience. We found that fCORT was negatively associated with growth rate in daughters, while it showed a positive association in sons. Lower offspring fCORT was associated with higher maternal baseline plasma CORT, and fCORT was higher in larger clutches and in those hatching later. The RTL of daughters was negatively associated with maternal RTL, whereas that of males was nearly independent of maternal RTL. Higher fCORT in ovo was associated with longer RTL at hatching in both sexes. Duckling body condition was mainly explained by egg weight, and sons had a slightly lower body condition. Our correlational results suggest that maternal effects may have heterogeneous and even diametrically opposed effects between the sexes during early development. Our findings also challenge the view that prenatal CORT exposure is invariably associated with shorter telomeres.

Investigation of the 2018 thick-billed murre (Uria lomvia) die-off on St. Lawrence Island rules out food shortage as the cause.

Die-offs of seabirds in Alaska have occurred with increased frequency since 2015. In 2018, on St. Lawrence Island, seabirds were reported washing up dead on beaches starting in late May, peaking in June, and continuing until early August. The cause of death was documented to be starvation, leading to the conclusion that a severe food shortage was to blame. We use physiology and colony-based observations to examine whether food shortage is a sufficient explanation for the die-off, or if evidence indicates an alternative cause of starvation such as disease. Specifically, we address what species were most affected, the timing of possible food shortages, and food shortage severity in a historical context. We found that thick-billed murres (Uria lomvia) were most affected by the die-off, making up 61% of all bird carcasses encountered during beach surveys. Thick-billed murre carcasses were proportionately more numerous (26:1) than would be expected based on ratios of thick-billed murres to co-occurring common murres (U. aalge) observed on breeding study plots (7:1). Concentrations of the stress hormone corticosterone, a reliable physiological indicator of nutritional stress, in thick-billed murre feathers grown in the fall indicate that foraging conditions in the northern Bering Sea were poor in the fall of 2017 and comparable in severity to those experienced by murres during the 1976-1977 Bering Sea regime shift. Concentrations of corticosterone in feathers grown during the pre-breeding molt indicate that foraging conditions in late winter 2018 were similar to previous years. The 2018 murre egg harvest in the village of Savoonga (on St. Lawrence Is.) was one-fifth the 1993-2012 average, and residents observed that fewer birds laid eggs in 2018. Exposure of thick-billed murres to nutritional stress in August, however, was no different in 2018 compared to 2016, 2017, and 2019, and was comparable to levels observed on St. George Island in 2003-2017. Prey abundance, measured by the National Oceanic and Atmospheric Administration in bottom-trawl surveys, was also similar in 2018 to 2017 and 2019, supporting the evidence that food was not scarce in the summer of 2018 in the vicinity of St. Lawrence Island. Of two moribund thick-billed murres collected at the end of the mortality event, one tested positive for a novel re-assortment H10 strain of avian influenza with Eurasian components, likely contracted during the non-breeding season. It is not currently known how widely spread infection of murres with the novel virus was, thus insufficient evidence exists to attribute the die-off to an outbreak of avian influenza. We conclude that food shortage alone is not an adequate explanation for the mortality of thick-billed murres in 2018, and highlight the importance of rapid response to mortality events in order to document alternative or confounding causes of mortality.