Intronic variation at the CHD1‐Z gene in Black‐tailed Godwits Limosa limosa limosa: correlations with fitness components revisited

Recently, Schroeder et al. (2010, Ibis 152: 368–377) suggested that intronic variation in the CHD1‐Z gene of Black‐tailed Godwits breeding in southwest Friesland, The Netherlands, correlated with fitness components. Here we re‐examine this surprising result using an expanded dataset (2088 birds sampled from 2004 to 2010 vs. 284 birds from 2004 to 2007). We find that the presence of the Z* allele (9% of the birds) is not associated with breeding habitat type, egg size, adult survival, adult body mass or adult body condition. The results presented here, when used in synergy with the previously reported results by Schroeder et al., suggest that there might be a tendency towards female adults with the Z* allele laying earlier clutches than adult females without the Z* allele. The occurrence of the Z* allele was also associated with a higher chick body mass and return rate. Chicks with the Z* allele that had hatched early in the breeding season were heavier at birth than chicks without the Z* allele and chicks with the Z* allele that had hatched late. Collectively, the results suggest that variation in the CHD1‐Z gene may indeed have arisen as a byproduct of selection acting on females during the egg fase and on chicks during the rearing stages of the reproductive cycle.     

Breeding Yellow-legged Gulls increase consumption of terrestrial prey after landfill closure

The ecology of opportunistic foragers can be highly dependent on anthropogenic food sources, such as landfills, resulting in changes in several ecological and demographic aspects. The total closure of several landfill sites and the use of deterrence systems to prevent access to the remaining open landfill sites in a region in the northern Iberian Peninsula provided an excellent opportunity to evaluate the consequences of landfills on the trophic ecology of an opportunistic forager, the Yellow-legged Gull Larus michahellis, using these resources. Based on stable isotope analyses, we used mixing models to estimate changes in diet before and after the closure of the majority of landfills in the breeding and the non-breeding season. We found a decrease in the consumption of food from landfills after their closure, which was accompanied by an increase in feeding on terrestrial prey (mostly earthworms), but only in the breeding season. Interestingly, we observed no increase in marine prey consumption after the landfill closures. In winter there was a decrease in terrestrial prey consumption, whereas the consumption of marine and, despite their reduce availability, landfill resources increased. Thus, and unlike when all the landfills were open, we detected significant trophic differences between breeding and non-breeding seasons. Additionally, we found significant trophic differences among colonies that were quite close to each other, but not between breeding adults and chicks. In conclusion, landfill closure or the use of deterrence systems had a direct impact on the trophic ecology of Yellow-legged Gulls; loss of refuse was mainly compensated for by prey of terrestrial origin when breeding, but not in winter. Thus, this species may experience foraging constraints in winter with potential effects on other life-history aspects including their dispersal, breeding and survival that needs further evaluation.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.     

Seasonal variation in pre‐fledging survival of lesser scaup Aythya affinis: hatch date effects depend on maternal body mass

Among temperate‐breeding birds, offspring survival and reproductive success are often inversely related to timing of breeding. The mechanisms that produce seasonal declines in offspring survival are not fully understood but may be related to temporal changes in parental quality, environmental quality, or both. We analyzed data for lesser scaup Aythya affinis to evaluate hypothesized effects of parental quality and date on pre‐fledging survival. Maternal quality, as indexed by body mass, did not have an independent effect on offspring survival in this species. Maternal body mass did not decline seasonally and did not have an independent effect on duckling survival. Although we did not detect an independent effect of hatch date on duckling survival, duckling survival declined seasonally for broods raised by lightweight females, indicating an interactive effect of maternal mass and date. We hypothesize that this interaction may be driven by seasonally declining food resources coupled with the influence of female condition on the ability to monopolize food resources or remain attentive to the brood. We also tested morphological predictions of the date hypothesis by examining physical characteristics of ducklings. When corrected for age and size, late‐hatched ducklings tended to have marginally larger digestive systems and smaller leg muscles than did early‐hatched birds. Abundances of intestinal parasites acquired through diet decreased marginally in late‐hatched ducklings. Results for digestive system and parasite infection patterns suggested that later‐hatched broods may shift diets, consistent with a contribution of environmental factors to seasonal variation in offspring survival. Taken together, our results suggest that both female attributes and environmental conditions may influence seasonal patterns of offspring survival in this species.     

Effect of neck collars on the body condition of migrating Greater Snow Geese

Markers are widely used to study behavior, migration, and the life history traits of birds such as survival, dispersal, and reproductive success. The presence of neck collars has been shown to impact the breeding propensity of adult female Greater Snow Geese (Chen caerulescens atlantica), but not their survival rates. We evaluated the hypothesis that the reduction in breeding propensity in neck‐collared birds was due to a reduction in the body condition of these long‐distance migrants that rely on a partial capital breeding strategy. Our study was conducted during 4 consecutive years along the St. Lawrence estuary in Quebec, Canada, a major spring staging area for these geese. We captured and marked 2552 geese with collars and 34 were recaptured in subsequent years at the same site. After controlling for confounding variables such as year and date of capture, we found that the presence of a neck collar reduced body condition of females during spring staging. Female Greater Snow Geese lost an average of 105.5 ± 39.1 (SE) g (4% of body mass) after carrying a collar for 1 yr and an average of 81.9 ± 43.6 g compared to original mass when recaptured 2 or 3 yr later. Our results suggest that the previously reported reduction in breeding propensity of neck‐collared geese may be due to a reduction in body condition during spring staging. Neck collars could negatively affect the body condition of female Greater Snow Geese by increasing their energy expenditure (due to increased drag during flight or to chronic stress) or reducing their foraging efficiency.     

Nonbreeder birds at colonies display qualitatively similar seasonal mass change patterns as breeders

The difficulty in studying nonbreeding birds means that little is known about them or their resource requirements, despite forming a large and significant component of a population. One way to assess food requirements is to examine changes in body mass, because it indicates the amount of food acquired. In terms of body mass changes, our expectation is that nonbreeders will either (a) be in poorer condition than the breeders which potentially explains why they do not breed or (b) remain at a stable higher mass as they are unconstrained by the physiological costs associated with rearing chicks. Here, we interrogate body mass datasets of breeding and nonbreeding birds of two penguin species to assess these predictions and determine whether differences in mass exist between these two groups throughout the breeding season. The first dataset is from a wild Adélie penguin population, where bird mass was recorded automatically and breeding status determined from a resighting program. A second population of captive gentoo penguins were weighed regularly each breeding season. We demonstrate that although there were times in each year when breeders were heavier than their nonbreeding counterparts for both populations, the mass changes showed qualitatively similar patterns throughout the breeding season irrespective of breeding status. Heavier breeders at times during the breeding season are not unexpected but the overall similar pattern of mass change irrespective of breeding status is in contrast to expectations. It appears that breeding status per se and the constraints that breeding places on birds are not the only driver of changes in mass throughout the breeding season and, although not explicitly studied here, the role of hormones in driving changes in appetite could be key to explain these results. These results present a significant step toward understanding food requirements of nonbreeders in avian populations.     

Sex‐specific mass loss in chick‐rearing South Polar Skuas Stercorarius maccormicki– stress induced or adaptive?

Mass loss of chick‐rearing birds can be the direct consequence of physiological stress (reproductive stress hypothesis) or an adaptive mass adjustment in response to the increased demands on flight efficiency during the flight‐intensive chick‐rearing period (adaptive mass loss hypothesis). To test which of these hypotheses best explains mass loss in South Polar Skuas Stercorarius maccormicki rearing chicks, a food supplementation experiment was carried out in the austral summer 2000/01 at King George Island, Antarctica. Half of the breeding pairs were fed about 20% of the chick's daily energy demand every second day and chick growth and adult nest attendance were recorded. Parents were caught at the start and the end of chick‐rearing to calculate adult mass loss. Male parents of food‐supplemented pairs attended their nest territories more than control males but females kept their attendance constant. Chick growth was only minimally affected and the treatment probably had no fitness consequences. Male Skuas in control pairs had a higher deviation from the body size–mass regression at the end of chick‐rearing compared with the start, supporting the stress hypothesis, whereas female deviation remained unchanged. Males of food‐supplemented pairs were heavier than unsupplemented males at the end of the breeding cycle but not significantly so. Food‐supplemented females were lighter at the end, supporting the adaptive mass loss hypothesis. Adult mass loss is thus best explained by the reproductive stress hypothesis in males but by the adaptive mass loss hypothesis in females. However, the two hypotheses are not mutually exclusive and the results do not exclude the possibility that mass loss in females is stress‐induced but the amount of mass lost is an adaptive adjustment to the reliability of the food supply. The finding that members of a breeding pair may follow different strategies of mass adjustment has implications for the use of mass loss as an index of parental effort. Without knowing which strategy each sex has adopted it is of little use to compare mass loss between parents.     

Variation in selection pressure acting on body size by age and sex in a reverse sexual size dimorphic raptor

Body size strongly influences fitness, with larger individuals benefiting in terms of both greater productivity and survivorship; for reverse sexual size dimorphic (RSD) species, this relationship may be more complex. We examined the selection pressures acting on body size in male and female Merlins Falco columbarius to assess whether larger or smaller individuals of this RSD species were favoured in terms of survival and breeding performance. For males and females there were clear links between body size and survival but the exact relationship varied by sex. Among males, birds that survived each year class were larger than those that died and yearlings were on average smaller than older birds, but there were no measurable differences among adult males (age 2+). Among females, larger individuals aged 1 and 2 years were more likely to survive, but this size‐based pattern was not apparent in older age classes. Size early in life predicted the lifespan in male Merlins but not as strongly as for females and not for the largest individuals. Reproductive performance based on brood size was not associated with body size in either males or females, but there was a weak positive relationship between female body size and lifetime reproductive success. Selection appears to favour larger males and females but there is no indication that the population is evolving towards bigger individuals, perhaps in part due to selection against the largest birds. Increased survival may allow larger and higher quality individuals to occupy higher quality territories as they age and thereby to accrue greater lifetime reproductive success in the process.     

Cross‐fostering eggs reveals that female collared flycatchers adjust clutch sex ratios according to parental ability to invest in offspring

Across animal taxa, reproductive success is generally more variable and more strongly dependent upon body condition for males than for females; in such cases, parents able to produce offspring in above‐average condition are predicted to produce sons, whereas parents unable to produce offspring in good condition should produce daughters. We tested this hypothesis in the collared flycatcher (Ficedula albicollis) by cross‐fostering eggs among nests and using the condition of foster young that parents raised to fledging as a functional measure of their ability to produce fit offspring. As predicted, females raising heavier‐than‐average foster fledglings with their social mate initially produced male‐biased primary sex ratios, whereas those raising lighter‐than‐average foster fledglings produced female‐biased primary sex ratios. Females also produced male‐biased clutches when mated to males with large secondary sexual characters (wing patches), and tended to produce male‐biased clutches earlier within breeding seasons relative to females breeding later. However, females did not adjust the sex of individuals within their clutches; sex was distributed randomly with respect to egg size, laying order and paternity. Future research investigating the proximate mechanisms linking ecological contexts and the quality of offspring parents are able to produce with primary sex‐ratio variation could provide fundamental insight into the evolution of context‐dependent sex‐ratio adjustment.     

Influences of Parental Care and Food Deprivation on Regulation of Body Mass in a Burying Beetle

Changes in adult body mass during breeding can reveal how parents prepare energetically for care, the stress of care, and the need to terminate care in a state conducive for future reproduction. Interpreting changes in parent mass can be difficult, however, because temporal variation in body mass may reflect a constraint imposed by the stress of care, revealing conflict within the family, or a shift to a new body mass optimum adaptive for a different stage of the breeding cycle. Here, we examined the effect of food deprivation and parenting on variation in female body mass of Nicrophorus orbicollis, an insect in which parents and offspring share a common food resource (a prepared carrion ball). Female parents demonstrated a remarkable degree of regulation of body mass: Despite varied periods of food deprivation (0–8 d) prior to discovery of a carcass, females attained a similar body mass (108.3–109.2% of pre‐deprivation mass) at the time of larval hatching. Females attained a greater body mass in anticipation of rearing a greater number of young. Mothers lost mass during active parental care, and mass at the end of caregiving was less in mothers that reared more and heavier young. Body mass at the end of care was less than the preferred mass for females searching for a carcass, indicating that the mothers sacrificed self‐maintenance and future reproductive potential for their current brood. Contrary to prediction, pre‐breeding food deprivation had no effect on offspring size or on female condition at the end of care. We conclude that there is a limited degree of conflict over the sharing of food among N. orbicollis parents and offspring, but that this conflict is not exacerbated by food deprivation prior to breeding.     

Short‐term changes in body condition in relation to habitat and rainfall abundance in American redstarts Setophaga ruticilla during the non‐breeding season

In migratory birds, environmental conditions during the stationary period of the non‐breeding season are crucial to consider because they ultimately affect the fitness of individuals by influencing their subsequent migration, breeding success and survival. Although a few studies have investigated the influence of non‐breeding habitat on the capacity of individuals to cope with long‐term seasonal rainfall fluctuations, it remains unknown how habitat quality and variations in rainfall abundance–at a monthly scale–interact to affect non‐breeding condition of migrating birds. In this study, we examined the influence of monthly changes in rainfall abundance on body condition of non‐breeding female redstarts Setophaga ruticilla living either in a high quality habitat (mangrove) or in a low quality habitat (scrub). Body condition of both mangrove and scrub redstarts showed important variations over the study period, demonstrating for the first time that body condition of non‐breeding female redstarts can change rapidly in response to short‐term fluctuations in rainfall. Importantly, we found that female redstarts living in mangrove were usually in better condition during periods of low rainfall compared to females living in scrub. However, body condition did not differ between mangrove females and scrub females during an episode of frequent, heavy precipitation. Importantly, our study also demonstrated that the duration of a perturbation is an important determinant of body condition in redstarts since a prolonged drought resulted in similar low body condition for birds from both habitats. Age was not correlated with body condition whatever the habitat and the rainfall conditions. Our results demonstrate that high quality habitat can temporarily reduce the deleterious effect of a short‐term drought on body condition, but also, that a habitat of low quality does not constrain individuals when climatic conditions are optimal.     

Carry‐over effects provide linkages across the annual cycle of a Neotropical migratory bird,the Louisiana Waterthrush Parkesia motacilla

Population limitation models of migratory birds have sought to include impacts from events across the full annual cycle. Previous work has shown that events occurring in winter result in some individuals transitioning to the breeding grounds earlier or in better physical condition than others, thereby affecting reproductive success (carry‐over effects). However, evidence for carry‐over effects from breeding to wintering grounds has been shown less often. We used feather corticosterone (CORT_f) levels of the migratory Louisiana Waterthrush Parkesia motacilla as a measure of the physiological state of birds at the time of moult on the breeding territory to investigate whether carry‐over effects provide linkages across the annual cycle of this stream‐obligate bird. We show that birds arriving on wintering grounds with lower CORT_f scores, indicating reduced energetic challenges or stressors at the time of moult, occupied higher quality territories, and that these birds then achieved a better body condition during the overwinter period. Body condition, in turn, was important in determining whether adult birds returned the following winter, with birds in better condition returning at higher rates. Together these data suggest a carry‐over effect from the breeding grounds to the wintering grounds that is further extended with respect to annual return rates. Very few other studies have linked conditions during the previous breeding season with latent effects during the subsequent overwintering period or with annual survival. This study shows that the effects of variation in energetic challenges or stressors can potentially carry over from the natal stream and accumulate over more than one life‐history period before being manifested in reduced survival. This is of particular relevance to models of population limitation in migratory birds.     

Sex allocation and sex‐specific parental investment in an endangered seabird

Biased offspring sex ratio is relatively rare in birds and sex allocation can vary with environmental conditions, with the larger and more costly sex, which can be either the male or female depending on species, favoured during high food availability. Sex‐specific parental investment may lead to biased mortality and, coupled with unequal production of one sex, may result in biased adult sex ratio, with potential grave consequences on population stability. The African Penguin Spheniscus demersus, endemic to southern Africa, is an endangered monogamous seabird with bi‐parental care. Female adult African Penguins are smaller, have a higher foraging effort when breeding and higher mortality compared with adult males. In 2015, a year in which environmental conditions were favourable for breeding, African Penguin chick production on Bird Island, Algoa Bay, South Africa, was skewed towards males (1.5 males to 1 female). Males also had higher growth rates and fledging mass than females, with potentially higher post‐fledging survival. Female, but not male, parents had higher foraging effort and lower body condition with increasing number of male chicks in their brood, thereby revealing flexibility in their parental strategy, but also the costs of their investment in their current brood. The combination of male‐biased chick production and higher female mortality, possibly at the juvenile stage as a result of lower parental investment in female chicks, and/or at the adult stage as a result of higher parental investment, may contribute to a biased adult sex ratio (ASR) in this species. While further research during years of contrasting food availability is needed to confirm this trend, populations with male‐skewed ASRs have higher extinction risks and conservation strategies aiming to benefit female African Penguin might need to be developed.     

Variation in the adult body mass of Wilson’s storm petrels Oceanites oceanicus during breeding

Mass loss of breeding birds might be due to the physiological stress of breeding or it could be an adaptation to lower the costs of flight to the feeding areas. We examined the natural variation in the adult body mass of Wilson’s storm petrels Oceanites oceanicus on King George Island, South Shetland Islands over four breeding seasons. During two seasons, the prey abundance was high, while it was poor during the other two seasons. Only breeding birds were sampled; the fluctuations in mass were similar among males and females. During incubation, the mass of the adults was high in good seasons and low in poor seasons. Thus, body mass during incubation was determined by energetic constraints. However, during chick feeding, adults lost mass in the good seasons but gained mass in the poor seasons, suggesting that mass loss during chick rearing is not primarily caused by stress, but is regulated adaptively. Adults in poor conditions may buffer against unpredictable food supply by increasing their own body mass, even at the expense of the chick. Reduced body condition at the beginning of the breeding season was associated with reduced egg volumes and late laying, suggesting that the initial body condition influenced the level of investment in the current breeding attempt.     

Seasonal variation in food availability influences the breeding strategy of White‐collared Blackbirds Turdus albocinctus on the Tibetan Plateau

Parent birds show a continuous spectrum of breeding strategies, ranging from a low‐fecundity and high‐survival pattern to a high‐fecundity, low‐survival pattern. Investigations of parental breeding strategies under variable environmental conditions can illustrate how parents trade‐off the benefits and costs of these two extreme strategies. White‐collared Blackbirds Turdus albocinctus can breed twice a year on the Tibetan Plateau. We show that both life‐history traits and parental feeding behaviour differ between these two breeding attempts. In the first attempt, the birds produced small clutches and fledged a small number of nestlings of high body condition. In the second attempt, they produced larger clutches and fledged more nestlings of lower body condition. Males made greater contributions to brood provisioning compared with females in the first attempt but there was no sex difference in brood provisioning in the second attempt. In the first attempt, producing smaller clutches can shorten the nestling period, and the increased male contribution to brood provisioning can protect the energy reserves of females. Thus, females can begin a second attempt sooner and produce larger clutches. During the second nesting attempt, when conditions are warmer and wetter, parents rely on a broader array of food types (both invertebrates and plant material, primarily berries) than during the first attempt, which includes only animal food such as arthropods and annelids. We suggest that this difference in breeding strategies between nesting attempts and sexes is in part influenced by marked seasonal variation in food availability.     

Breeding limits foraging time: evidence of interrupted foraging response from body mass variation in a tropical environment

Birds should store body reserves if starvation risk is anticipated; this is known as an ‘interrupted foraging response’. If foraging remains unrestricted, however, body mass should remain low to limit the predation risk that gaining and carrying body reserves entails. In temperate environments mass gain in female birds during breeding is often attributed to egg formation and mass loss after incubation to flight adaptation or the effect of reproductive workload, rather than as a result of an adaptive interrupted foraging response to the limited foraging time or unpredictable foraging conditions that breeding demands. In tropical environments, foraging conditions vary more within the breeding season than in temperate environments, and so studies in tropical environments are more suited to decouple the potentially confounded effects of increase in body reserves versus egg formation on the body mass of breeding birds. In this study, we test whether breeding results in an interrupted foraging response in a tropical savannah system using body mass data collected over a 15‐year period from female common bulbuls Pycnonotus barbatus. This species breeds both in the wet and dry season, despite fewer resources being available in the dry season. Breeding stage predicted female body mass: body mass peaked abruptly during incubation, but was not closely associated with the egg‐laying stage, and declined during brood rearing. Breeding females were heavier in the dry season than in the wet season. In the dry season, heavier birds were more likely to incubate eggs or brood chicks. These observations suggest that increased body reserves are required to buffer the consequence of limited foraging time or impoverished foraging conditions, which may be most pronounced during incubation and in the dry season, respectively. Such mass increases are consistent with an interrupted foraging response, which may apply to temperate zone birds experiencing foraging restrictions during breeding.     

The timing, duration and pattern of moult and its relationship to breeding in a population of the European Greenfinch Carduelis chloris

Moult was studied in 1 year among Greenfinches trapped in a garden in east‐central England. Over the period June–December 2003, 333 captures of 179 individual adults provided information on breeding condition, moult, body weight, sex and age (yearling or older adult, equivalent to birds in their second or later calendar years, respectively). About 95% of all birds (sex and age groups combined) started primary feather moult from 2 July to 14 August, and finished from 10 October to 22 November. The mean date of moult onset in the population as a whole was 24 July. On average, males began 8 days before females, and yearlings began 6 days before older birds. The mean duration of moult was 100 days, whether the figure was calculated for the population as a whole or just for the 36 individual birds that were caught more than once during moult. However, moult rate was slightly slower, and moult duration slightly longer, in yearlings than in older adults of both sexes. No evidence was found for any systematic relationship between moult onset date and rate (duration). Breeding and moult overlapped by up to 5 weeks or more in individual birds, and some birds probably started to moult as early as the incubation stage of their last clutch of the season. The cloacal protuberance (taken as indicative of breeding condition) had regressed in all males by the time the fifth primary was shed, and the brood patch had regressed and re‐feathered in all females by the time the fourth primary was shed. The bulk of feather replacement in the secondary, tail and body tracts occurred in the second half of primary moult, and after cloacal protuberances and brood patches were completely regressed. In all birds examined near the end of primary moult the secondaries were still growing, and would have continued growth for up to another 19 days or more, extending the end of the moulting season into December. Body mass during moult was affected significantly by sex and age, as well as by time of day, amount of food in gullet, reproductive condition and date. No firm evidence emerged that body mass was affected by moult stage, after allowing for effects of date and other variables (although there was a non‐significant negative relationship between moult stage and body mass in males). In the population as a whole, the breeding season (from first egg‐laying to independence of last young) was spread over 21 weeks and moult over 24 weeks. With an overlap between the two events at the population level of up to 9 weeks, the two processes together took up to 36 weeks, some 69% of the year.     

Central–periphery gradient of individual quality within a colony of Black‐headed Gulls

Central nesting sites within avian colonies are often more profitable in terms of fitness, as they can offer better protection against predators compared with colony edges. Thus, central sites are expected to attract high‐quality individuals, which should produce a clear central–periphery gradient in the phenotypic quality of nesting individuals and reproductive output within colonies. The aim of this study was to assess spatial patterns of individual quality within a colony of a common larid, the Black‐headed Gull Chroicocephalus ridibundus. We found that laying was advanced in the central zone of the colony when compared with the periphery, and central pairs tended to lay larger eggs and had higher hatching success than peripheral pairs. Centrally nesting individuals had higher blood haemoglobin concentrations, size‐corrected body mass and heterozygosity (males only), indicating their better condition and higher genetic quality. In contrast, central females had lower plasma albumin concentration, which could possibly be attributed to their greater investment in egg production. Finally, pairs breeding in the central zone of the colony had an elevated heterophil/lymphocyte ratio, suggesting that social stress can be recognized as an important cost of nesting in dense aggregations. The results provide empirical support for central–periphery gradients in the individual quality of colonially nesting Black‐headed Gulls, indicating that spatial gradients in pair quality should be carefully taken into account when conducting research on colonial birds.     

Body condition dynamics and the cost‐of‐delay hypothesis in a temperate‐breeding duck

Pre‐breeding body condition is an important determinant of reproductive success in birds, largely through its influence on timing of breeding. Declines in clutch size and recruitment probability within breeding seasons indicate a tradeoff may exist between the number of young (clutch size) and quality of young (recruitment probability). We explored local drivers of pre‐breeding body condition and tested predictions of the cost‐of‐delay hypothesis in female lesser scaup Aythya affinis. Yearling females arrived on the study site in lower body condition than older females, but both age classes had similar rates of body condition gain on the breeding grounds prior to nesting. Rates of body condition gain were positively influenced by water temperature, a proxy for wetland phenology. The effect of water level was asymptotic and interacted with water temperature, with greater rates of gain in body condition occurring in years with low water levels. Our results supported the predicted response of clutch size to the rate of pre‐breeding body condition gain. After accounting for lay date, clutch size was positively related to the rate of body condition gain (= 0.08 ± 0.039). We did not find support for a predicted interaction between rate of body condition gain and intra‐seasonal decline in clutch size (= 0.01 ± 0.01). Our results indicate that local conditions during pre‐breeding influence body condition dynamics in female lesser scaup, which subsequently affects clutch size.