Energy supply during nocturnal endurance flight of migrant birds: effect of energy stores and flight behaviour.

BACKGROUND: Migrating birds fly non-stop for hours or even for days. They rely mainly on fat as fuel complemented by a certain amount of protein. Studies on homing pigeons and birds flying in a wind-tunnel suggest that the shares of fat and protein on total energy expenditure vary with flight duration and body fat stores. Also, flight behaviour, such as descending flight, is expected to affect metabolism. However, studies on free flying migrant birds under natural conditions are lacking. METHODS: On a Swiss Alpine pass, we caught three species of nocturnal migrant passerines out of their natural migratory flight. Since most night migrants start soon after dusk, we used time since dusk as a measure of flight duration. We used plasma concentrations of metabolites of the fat, protein, and carbohydrate metabolism as indicators of relative fuel use. We used flight altitudes of birds tracked with radar and with atmospheric pressure loggers to characterize flight behaviour. RESULTS: The indicators of fat catabolism (triglycerides, very low-density lipoproteins, glycerol) were positively correlated with body energy stores, supporting earlier findings that birds with high fat stores have a higher fat catabolism. As expected, plasma levels of triglycerides, very low-density lipoproteins, glycerol and ß-hydroxy-butyrate increased at the beginning of the night, indicating that nocturnal migrants increased their fat metabolism directly after take-off. Surprisingly, fat catabolism as well as glucose levels decreased in the second half of the night. Data from radar observations showed that the number of birds aloft, their mean height above ground and vertical flight speed decreased after midnight. Together with the findings from atmospheric pressure-loggers put on three species, this shows that nocturnal migrants migrating over continental Europe descend slowly during about 1.5 h before final landfall at night, which results in 11-30% energy savings according to current flight models. CONCLUSIONS: We suggest that this slow descent reduces energy demands to an extent which is noticeable in the plasma concentration of lipid, protein, and carbohydrate metabolites. The slow descent may facilitate the search for a suitable resting habitat and serve to refill glycogen stores needed for foraging and predator escape when landed.

How woodcocks produce the most brilliant white plumage patches among the birds.

Until recently, and when compared with diurnal birds that use contrasting plumage patches and complex feather structures to convey visual information, communication in nocturnal and crepuscular species was considered to follow acoustic and chemical channels. However, many birds that are active in low-light environments have evolved intensely white plumage patches within otherwise inconspicuous plumages. We used spectrophotometry, electron microscopy, and optical modelling to explain the mechanisms producing bright white tail feather tips of the Eurasian woodcock Scolopax rusticola. Their diffuse reflectance was approximately 30% higher than any previously measured feather. This intense reflectance is the result of incoherent light scattering from a disordered nanostructure composed of keratin and air within the barb rami. In addition, the flattening, thickening and arrangement of those barbs create a Venetian-blind-like macrostructure that enhances the surface area for light reflection. We suggest that the woodcocks have evolved these bright white feather patches for long-range visual communication in dimly lit environments.

Exogenous corticosterone and melanin-based coloration explain variation in juvenile dispersal behaviour in the barn owl (Tyto alba).

Natal dispersal affects many processes such as population dynamics. So far, most studies have examined the intrinsic and extrinsic factors that determine the distance between the place of birth and of first breeding. In contrast, few researchers followed the first steps of dispersal soon after fledging. To study this gap, we radio-tracked 95 barn owl nestlings (Tyto alba) to locate their diurnal roost sites from the fledging stage until December. This was used to test whether the age of nest departure, post-fledging movements and dispersal distance were related to melanin-based coloration, which is correlated to fitness-related traits, as well as to corticosterone, a hormone that mediates a number of life history trade-offs and the physiological and behavioural responses to stressful situations. We found that the artificial administration of corticosterone delayed the age when juveniles left their parental home-range in females but not in males. During the first few months after fledging, longer dispersal distances were reached by females compared to males, by individuals marked with larger black feather spots compared to individuals with smaller spots, by larger individuals and by those experimentally treated with corticosterone. We conclude that the onset and magnitude of dispersal is sensitive to the stress hormone corticosterone, melanin-based coloration and body size.

Factors modulating the behavioral and physiological stress responses: Do they modify the relationship between flight initiation distance and corticosterone reactivity?

Understanding how vulnerable species are to new stressors, such as anthropogenic changes, is crucial for mitigating their potential negative consequences. Many studies have investigated species sensitivity to human disturbance by focusing on single behavioral or physiological parameters, such as flight initiation distance and glucocorticoid levels. However, little is known about the differential effect that modulating factors might have on behavioral versus physiological stress responses across species. This lack of knowledge make difficult to understand the relationship between both types of reactions, and thus to assess to what extent a behavioral reaction is representative of an internal physiological stress response or vice versa. We collected published data on bird flight initiation distances (FID) and corticosterone (CORT) responses, the two most frequently used indicators of stress reaction. We then investigated how spatio-temporal factors or species-specific characteristics relate to these behavioral and physiological stress responses, and potentially modify the relationship between them. Additionally, we evaluated the strength of the correlation between the two stress responses (behavioral and physiological). Our findings showed that FID and CORT responses were poorly correlated across species, and the lack of correlation was attributable to modulating factors (e.g. latitude and body mass) which influence behavior and physiology differently. These modulating factors, therefore, should be taken into consideration to better interpret FID and CORT responses in the context of species vulnerability to stress.

Glucocorticoid levels are linked to lifetime reproductive success and survival of adult barn owls.

Glucocorticoid hormones, such as corticosterone, are crucial in regulating daily life metabolism and energy expenditure, as well as promoting short-term physiological and behavioural responses to unpredictable environmental challenges. Therefore, glucocorticoids are considered to mediate trade-offs between survival and reproduction. Relatively little is known about how selection has shaped glucocorticoid levels. We used 15 years of capture-recapture and dead recovery data combined with 13 years of corticosterone and breeding success data taken on breeding barn owls (Tyto alba) to investigate such trade-offs. We found that survival was positively correlated with stress-induced corticosterone levels in both sexes, whereas annual and lifetime reproductive success (i.e. the sum of young successfully fledged during the entire reproductive career) was positively correlated with both baseline and stress-induced corticosterone levels in females only. Our results suggest that, in the barn owl, the stress-induced corticosterone response is a good proxy for adult survival and lifetime reproductive success. However, selection pressure appears to act differently on corticosterone levels of males and females.

Determinants and constraints of feather growth.

During the periodic moult of the plumage of birds, a fast regrowth of feathers would shorten the time of reduced plumage functionality. However, it has long been known that feather growth-rate is limited and that long feathers take disproportionally longer to grow than small feathers, which has severe consequences on moult duration and the completeness of moult in large birds. The reasons for the limitations of feather-growth must be related to the size and/or functions of the feather follicle, but are largely unknown. Here we measured the size of the feather follicle (taking calamus width as a proxy) and related it to parameters of feather growth (feather growth-rate by mass and by length) and feather structure (feather length, mass, massiveness [mass of feather material per mm feather-length]). We used three independent datasets which allowed for interspecific analyses, and for intraspecific comparisons of differently structured feathers within the framework of biological scaling. We found that the cross-sectional area of the calamus (as a proxy of feather follicle size) was directly proportional to feather growth-rate by mass. Hence, factors acting at a two-dimensional scale (possibly nutrient supply to the growing feather) determines feather growth rate by mass, rather than the linear arrangement of stem cells (in a circular configuration) as had previously been assumed. Feather follicle size was correlated with both feather length and massiveness, hence it seems to be adapted to some extent to feather structure. Feather growth-rate by length was dependent on both the feather material produced per unit time (growth-rate by mass) and the amount of material deposited per unit feather-length. Follicle size not only determines feather growth-rate by mass, but also directly the structural design (shape, number of barbs, etc.) of a feather. Therefore, feather growth-rate is severely constrained by the requirements imposed by the structural feather design.

Human recreation decreases antibody titre in bird nestlings: an overlooked transgenerational effect of disturbance.

Outdoor recreational activities are booming and most animals perceive humans as predators, which triggers behavioural and/or physiological reactions [e.g. heart rate increase, activation of the hypothalamic-pituitary-adrenal (HPA) axis]. Physiological stress reactions have been shown to affect the immune system of an animal and therefore may also affect the amount of maternal antibodies a female transmits to her offspring. A few studies have revealed that the presence of predators affects the amount of maternal antibodies deposited into eggs of birds. In this study, using Eurasian blue and great tit offspring (Cyanistes caeruleus and Parus major) as model species, we experimentally tested whether human recreation induces changes in the amount of circulating antibodies in young nestlings and whether this effect is modulated by habitat and competition. Moreover, we investigated whether these variations in antibody titre in turn have an impact on hatching success and offspring growth. Nestlings of great tit females that had been disturbed by experimental human recreation during egg laying had lower antibody titres compared with control nestlings. Antibody titre of nestling blue tits showed a negative correlation with the presence of great tits, rather than with human disturbance. The hatching success was positively correlated with the average amount of antibodies in great tit nestlings, independent of the treatment. Antibody titre in the first days of life in both species was positively correlated with body mass, but this relationship disappeared at fledging and was independent of treatment. We suggest that human recreation may have caused a stress-driven activation of the HPA axis in breeding females, chronically increasing their circulating corticosterone, which is known to have an immunosuppressive function. Either, lower amounts of antibodies are transmitted to nestlings or impaired transfer mechanisms lead to lower amounts of immunoglobulins in the eggs. Human disturbance could, therefore, have negative effects on nestling survival at early life-stages, when nestlings are heavily reliant on maternal antibodies, and in turn lead to lower breeding success and parental fitness. This is a so far overlooked effect of disturbance on early life in birds.

Seasonal changes in yolk hormone concentrations carry-over to offspring traits.

Yolk hormones are substances which transmit non-genetic factors from the mother to the next generation. The systematic changes of yolk hormone concentrations within asynchronously hatching clutches have been interpreted as a means to adaptively shape the offspring's phenotype. However, in synchronously hatching clutches the role of yolk hormones is less understood. We investigated whether seasonal changes between eggs in the yolk hormones testosterone (Testo), progesterone (Prog) and corticosterone (Cort) also occur in the grey partridge, a synchronously hatching precocial species without direct food competition between siblings. Specifically we asked whether yolk hormone concentrations systematically vary with season and whether they affect the offspring's hatching mass, mass gain, circulating baseline and stress-induced Cort. Additionally, we investigated the effect of genetic background and food availability on yolk hormone concentrations by subjecting grey partridge hens of two strains (wild and domesticated) to two different feeding regimes (predictable vs. unpredictable feeding) during egg laying. We hypothesized that egg hormone concentrations change over the season, but breeding in captivity over many generations and ad libitum food access could have resulted in domestication effects which abolished potential seasonal effects. Results showed that progressing season had a strong positive effect on yolk Prog and yolk Testo, but not on yolk Cort. Feeding regimes and strain had no effect on yolk hormones. Offspring mass and mass gain increased and baseline Cort decreased with progressing season. In addition, yolk Testo correlated positively with offspring mass gain and negatively with baseline Cort, while yolk Prog had a positive correlation with baseline Cort. Strain and feeding regimes of the mother had no effect on offspring traits. In conclusion, grey partridge chicks hatching late in the season might benefit from the increased concentrations of the growth-stimulating yolk Testo and by this catch-up in development. Hence, yolk hormone concentration could adaptively shape the offspring phenotype in a precocial species.

Baseline and stress-induced corticosterone levels are heritable and genetically correlated in a barn owl population.

The hypothalamic-pituitary-adrenal (HPA) axis is responsible for the regulation of corticosterone, a hormone that is essential in the mediation of energy allocation and physiological stress. As a continuous source of challenge and stress for organisms, the environment has promoted the evolution of physiological adaptations and led to a great variation in corticosterone profiles within or among individuals, populations and species. In order to evolve via natural selection, corticosterone levels do not only depend on the strength of selection exerted on them, but also on the extent to which the regulation of corticosterone is heritable. Nevertheless, the heritability of corticosterone profiles in wild populations is still poorly understood. In this study, we estimated the heritability of baseline and stress-induced corticosterone levels in barn owl (Tyto alba) nestlings from 8 years of data, using a multivariate animal model based on a behavioural pedigree. We found that baseline and stress-induced corticosterone levels are strongly genetically correlated (r = 0.68-0.80) and that the heritability of stress-induced corticosterone levels (h2 = 0.24-0.33) was moderate and similar to the heritability of baseline corticosterone levels (h2 = 0.19-0.30). These findings suggest that the regulation of stress-induced corticosterone and baseline levels evolves at a similar pace when selection acts with the same intensity on both traits and that contrary to previous studies, the evolution of baseline and stress-induced level is interdependent in barn owls, as they may be strongly genetically correlated.

Acute and Chronic Lead Exposure in Four Avian Scavenger Species in Switzerland.

Despite irrefutable evidence of its negative impact on animal behaviour and physiology, lethal and sublethal lead poisoning of wildlife is still persistent and widespread. For scavenging birds, ingestion of ammunition, or fragments thereof, is the major exposure route. In this study, we examined the occurrence of lead in four avian scavengers of Switzerland and how it differs between species, regions, and age of the bird. We measured lead concentration in liver and bone of the two main alpine avian scavengers (golden eagle Aquila chrysaetos and bearded vulture Gypaetus barbatus) over the entire area of the Swiss Alps and two of the main avian scavengers occurring in the lowlands of Switzerland (red kite Milvus milvus and common raven Corvus corax). Of those four species, only the bearded vulture is an obligate scavenger. We found that lead burdens in the two alpine avian scavengers were higher than those found for the same species elsewhere in Europe or North America and reached levels compatible with acute poisoning, whereas lead burdens of the two lowland avian scavengers seemed to be lower. Several golden eagles, but only one red kite with abnormally high bone lead concentrations were found. In all four species, a substantial proportion of birds had elevated levels which presumably represent recent (liver lead levels) or past (bone lead levels) uptake of sublethal doses of lead.

Effect of human recreation on bird anti-predatory response.

Wildlife perceive humans as predators, and therefore normally flushes. Flight initiation distance (FID) is the distance a human can approach an animal at a steady pace until it flushes. Recently, several studies showed differences in within-species FID according to human presence by comparing urban and rural habitats, with urban birds showing reduced FIDs. However, urban and rural habitats also differ in structure, which might affect FID. Therefore, in order to understand the real effect of human presence, we investigated whether differences in FID are also present in natural habitats (forests), differing only in the intensity of human use for recreation. We found that human frequentation had a distinct effect on bird escape responses, with shorter FIDs in forests more-heavily frequented by humans than in forests rarely visited by humans. Whether this finding is driven by non-random spatial distribution of personalities (shy vs. bold) or phenotypic plasticity (habituation to humans) cannot be assessed with our data. Studies relying on FIDs should also incorporate human recreation intensity, as this affects the measurements strongly.

Experimental evidence of human recreational disturbance effects on bird-territory establishment.

The worldwide increase in human outdoor activities raises concerns for wildlife. Human disturbances, even at low levels, are likely to impact species during sensitive periods of the annual cycle. However, experimental studies during the putative sensitive period of territory establishment of birds which not only investigate low disturbance levels, but which also exclude the effect of habitat modification (e.g. walking trails) are lacking. Here, we experimentally disturbed birds in forest plots by walking through twice a day during territory establishment. Later we compared the breeding bird community of experimentally disturbed plots with that of undisturbed control plots. We discovered that the number of territories (-15.0%) and species richness (-15.2%) in disturbed plots were substantially reduced compared with control plots. Species most affected included those sensitive to human presence (assessed by flight-initiation distances), open-cup nesters and above-ground foragers. Long-distance migrants, however, were unaffected due to their arrival after experimental disturbance took place. These findings highlight how territory establishment is a sensitive period for birds, when even low levels of human recreation may be perceived as threatening, and alter settlement decisions. This can have important implications for the conservation of species, which might go unnoticed when focusing only on already established birds.

Happy to breed in the city? Urban food resources limit reproductive output in Western Jackdaws.

Urban areas expand worldwide, transforming landscapes and creating new challenging habitats. Some bird species, mainly omnivorous feeding on human waste and cavity nesters, commonly breed in these habitats and are, therefore, regarded as urban-adapted. Although urban areas may provide new nesting sites and abundant human waste, the low breeding success found in some of these species suggests that the poor protein content in human waste might limit breeding parameters. We investigated whether the breeding success of a cavity nester and omnivorous species commonly breeding in urban areas, the Western Jackdaw (Corvus monedula), depended on the availability of good-quality non-urban food. We approached the objective by combining a literature review and experiments in the field. With the literature review, we compared jackdaw populations in different habitats across Europe and found that clutch size and number of fledglings per pair decreased with distance to non-urban foraging grounds, even after controlling for the effect of colony size, latitude, and climate. In two experiments, we tested whether the breeding success of urban pairs could be increased by supplementing high-quality food, first only during egg formation and second also until chick fledging. Food supplementation during egg formation led to larger eggs and higher hatching success than in urban control nests, but this did not result in higher chick survival. However, when food supplementation was prolonged until fledging in the second experiment, we observed a significant increase of nestling survival. These findings highlight that research and management actions should not only focus on species displaced by urbanization, but also on "urban-adapted" species, as they might be suffering from a mismatch between availability of nesting sites in buildings and adequate non-urban food resources. In these cases, nest sites should be provided in or close to adequate food resources.

Determinants of uncertainty in wildlife responses to human disturbance.

Outdoor recreation is increasing in intensity and space. Areas previously inaccessible are now being visited by ever-growing numbers of people, which increases human-wildlife encounters across habitats. This has raised concern among researchers and conservationists as, even in non-aggressive encounters, animals often perceive humans as predators and mount physiological and behavioural responses that can have negative consequences. However, despite all the research in recent decades, not many general patterns have emerged, especially at the level of populations, and many studies have yielded seemingly contradictory or inconclusive results. We argue that this is partly due to incomplete knowledge of the number and complexity of factors that may modulate the responses of animals. Thus, we aim to provide a conceptual approach intended to highlight the reasons that make it difficult to detect general patterns. We present a comprehensive compilation of factors modulating animal responses to humans at increasing levels (from sensory detection and immediate behavioural and physiological reactions, to changes in fitness and population trends), which may help understanding the uncertainty in the patterns. We observed that there are many modulating factors, which can be categorized as reflecting characteristics of the recreational activity itself (e.g. intensity of human presence), of the animals concerned (e.g. age or antipredatory strategy), and of the spatio-temporal context (e.g. habitat or timing of the encounter). Some factors appear to have non-linear and complex effects, which, if not considered, may lead to erroneous conclusions. Finally, we conclude that the difficulty in finding general patterns will be amplified at higher levels (i.e. at the level of populations), since as we proceed from one level to the next, the number of potential modulating factors accumulates, adding noise and obscuring direct associations between recreation and wildlife. More comprehensive knowledge about which (and how) factors affect animal responses across levels will certainly improve future research design and interpretation, and thus, our understanding of human recreational impacts on wildlife.

Social huddling and physiological thermoregulation are related to melanism in the nocturnal barn owl.

Endothermic animals vary in their physiological ability to maintain a constant body temperature. Since melanin-based coloration is related to thermoregulation and energy homeostasis, we predict that dark and pale melanic individuals adopt different behaviours to regulate their body temperature. Young animals are particularly sensitive to a decrease in ambient temperature because their physiological system is not yet mature and growth may be traded-off against thermoregulation. To reduce energy loss, offspring huddle during periods of cold weather. We investigated in nestling barn owls (Tyto alba) whether body temperature, oxygen consumption and huddling were associated with melanin-based coloration. Isolated owlets displaying more black feather spots had a lower body temperature and consumed more oxygen than those with fewer black spots. This suggests that highly melanic individuals display a different thermoregulation strategy. This interpretation is also supported by the finding that, at relatively low ambient temperature, owlets displaying more black spots huddled more rapidly and more often than those displaying fewer spots. Assuming that spot number is associated with the ability to thermoregulate not only in Swiss barn owls but also in other Tytonidae, our results could explain geographic variation in the degree of melanism. Indeed, in the northern hemisphere, barn owls and allies are less spotted polewards than close to the equator, and in the northern American continent, barn owls are also less spotted in colder regions. If melanic spots themselves helped thermoregulation, we would have expected the opposite results. We therefore suggest that some melanogenic genes pleiotropically regulate thermoregulatory processes.

Agricultural land use and human presence around breeding sites increase stress-hormone levels and decrease body mass in barn owl nestlings.

Human activities can have a suite of positive and negative effects on animals and thus can affect various life history parameters. Human presence and agricultural practice can be perceived as stressors to which animals react with the secretion of glucocorticoids. The acute short-term secretion of glucocorticoids is considered beneficial and helps an animal to redirect energy and behaviour to cope with a critical situation. However, a long-term increase of glucocorticoids can impair e.g. growth and immune functions. We investigated how nestling barn owls (Tyto alba) are affected by the surrounding landscape and by human activities around their nest sites. We studied these effects on two response levels: (a) the physiological level of the hypothalamus-pituitary-adrenal axis, represented by baseline concentrations of corticosterone and the concentration attained by a standardized stressor; (b) fitness parameters: growth of the nestlings and breeding performance. Nestlings growing up in intensively cultivated areas showed increased baseline corticosterone levels late in the season and had an increased corticosterone release after a stressful event, while their body mass was decreased. Nestlings experiencing frequent anthropogenic disturbance had elevated baseline corticosterone levels, an increased corticosterone stress response and a lower body mass. Finally, breeding performance was better in structurally more diverse landscapes. In conclusion, anthropogenic disturbance affects offspring quality rather than quantity, whereas agricultural practices affect both life history traits.

Distinct responses of baseline and stress-induced corticosterone levels to genetic and environmental factors.

Glucocorticoid (GC) hormones, i.e. corticosterone (CORT) in birds, support physiological homeostasis and facilitate adaptations to stressful situations. However, maintaining high GC levels are energetically costly and interfere with other physiological processes. To keep the balance of costs and benefits of GC hormones, various mechanisms act to adapt GC levels to environmental conditions on different timescales, i.e. over generations, between parents and their offspring and within the life-time of a single individual. We elucidated whether two strains (domesticated and wild) of grey partridges (Perdix perdix) differed in the developmental trajectories of baseline and stress response CORT throughout the first 80 days of life. We also explored the potential of prenatal and postnatal factors, e.g. parental origin, predictable vs. unpredictable food treatments, individual and social factors to modify these trajectories. Baseline CORT was similar between strains and unaffected by perinatal food treatments. It was negatively related to body size and body condition. Conversely, the CORT stress response was not markedly affected by physiological condition. It was stronger in wild than in domesticated birds and it increased with age. Birds subjected to prenatal unpredictable food supply exhibited an accelerated development of the CORT stress response which could reflect an adaptive maternal effect. We conclude that the vital role of baseline CORT may allow little adaptive scope since changes can quickly become detrimental. In contrast, the CORT stress response may show considerable adaptive potential which might ultimately support homeostasis in a changing environment.

Tick-borne pathogens in ticks collected from breeding and migratory birds in Switzerland.

From 2007 to 2010, 4558 migrating and breeding birds of 71 species were caught and examined for ticks in Switzerland. A total of 1205 specimens were collected; all were Ixodes ricinus ticks except one Ixodes frontalis female, which was found on a common chaffinch (Fringilla coelebs) for the first time in Switzerland. Each tick was analysed individually for the presence of Borrelia spp., Rickettsia spp., Anaplasma phagocytophilum and tick-borne encephalitis virus (TBEV). Altogether, 11.4% of birds (22 species) were infested by ticks and 39.8% of them (15 species) were carrying infected ticks. Bird species belonging to the genus Turdus were the most frequently infested with ticks and they were also carrying the most frequently infected ticks. Each tick-borne pathogen for which we tested was identified within the sample of bird-feeding ticks: Borrelia spp. (19.5%) and Rickettsia helvetica (10.5%) were predominantly detected whereas A. phagocytophilum (2%), Rickettsia monacensis (0.4%) and TBEV (0.2%) were only sporadically detected. Among Borrelia infections, B. garinii and B. valaisiana were largely predominant followed by B. afzelii, B. bavariensis, B. miyamotoi and B. burgdorferi ss. Interestingly, Candidatus Neoehrlichia mikurensis was identified in a few ticks (3.3%), mainly from chaffinches. Our study emphasizes the role of birds in the natural cycle of tick-borne pathogens that are of human medical and veterinary relevance in Europe. According to infection detected in larvae feeding on birds we implicate the common blackbird (Turdus merula) and the tree pipit (Anthus trivialis) as reservoir hosts for Borrelia spp., Rickettsia spp. and A. phagocytophilum.

Oxidative stress in endurance flight: an unconsidered factor in bird migration.

Migrating birds perform extraordinary endurance flights, up to 200 h non-stop, at a very high metabolic rate and while fasting. Such an intense and prolonged physical activity is normally associated with an increased production of reactive oxygen and nitrogen species (RONS) and thus increased risk of oxidative stress. However, up to now it was unknown whether endurance flight evokes oxidative stress. We measured a marker of oxidative damage (protein carbonyls, PCs) and a marker of enzymatic antioxidant capacity (glutathione peroxidase, GPx) in the European robin (Erithacus rubecula), a nocturnal migrant, on its way to the non-breeding grounds. Both markers were significantly higher in European robins caught out of their nocturnal flight than in conspecifics caught during the day while resting. Independently of time of day, both markers showed higher concentrations in individuals with reduced flight muscles. Adults had higher GPx concentrations than first-year birds on their first migration. These results show for the first time that free-flying migrants experience oxidative stress during endurance flight and up-regulate one component of antioxidant capacity. We discuss that avoiding oxidative stress may be an overlooked factor shaping bird migration strategies, e.g. by disfavouring long non-stop flights and an extensive catabolism of the flight muscles.

Corticosterone shifts reproductive behaviour towards self-maintenance in the barn owl and is linked to melanin-based coloration in females.

Trade-offs between the benefits of current reproduction and the costs to future reproduction and survival are widely recognized. However, such trade-offs might only be detected when resources become limited to the point where investment in one activity jeopardizes investment in others. The resolution of the trade-off between reproduction and self-maintenance is mediated by hormones such as glucocorticoids which direct behaviour and physiology towards self-maintenance under stressful situations. We investigated this trade-off in male and female barn owls in relation to the degree of heritable melanin-based coloration, a trait that reflects the ability to cope with various sources of stress in nestlings. We increased circulating corticosterone in breeding adults by implanting a corticosterone-releasing-pellet, using birds implanted with a placebo-pellet as controls. In males, elevated corticosterone reduced the activity (i.e. reduced home-range size and distance covered within the home-range) independently of coloration, while we could not detect any effect on hunting efficiency. The effect of experimentally elevated corticosterone on female behaviour was correlated with their melanin-based coloration. Corticosterone (cort-) induced an increase in brooding behaviour in small-spotted females, while this hormone had no detectable effect in large-spotted females. Cort-females with small eumelanic spots showed the normal body-mass loss during the early nestling period, while large spotted cort-females did not lose body mass. This indicates that corticosterone induced a shift towards self-maintenance in males independently on their plumage, whereas in females this shift was observed only in large-spotted females.