Plasticity of moult and breeding schedules in migratory European Stonechats Saxicola rubicola

Timing is crucial in seasonal environments. Passerine birds typically use a combination of physiological mechanisms and environmental cues to ensure that breeding, moult and migration occur without major temporal overlap and under the most favourable conditions. However, late in the breeding season some individuals initiate additional clutches , whereas others initiate moult. Such alternative strategies are thought to reflect trade‐offs between reproductive benefits and timely investment in maintenance and survival. The degree of seasonal plasticity differs between species, depending on the mechanisms that govern their annual routine. Migrants are generally under pressure to complete breeding and moult before the autumn departure and often show little plasticity. We studied seasonal plasticity of breeding and moult schedules in the European Stonechat Saxicola rubicola. This species, an obligate short‐distance migrant in Central Europe, sometimes initiates late clutches after typically at least two earlier breeding attempts. Based on life‐history theory and on observations in captivity, which revealed photoperiodic regulation of breeding and moult, we predicted relatively little seasonal plasticity in Stonechats. We further predicted that reproductive gains of late breeders should be offset by reduced survival. These predictions were tested on long‐term field data, using Underhill–Zucchini models to estimate moult. Late breeding occurred in c. 40% of pairs and increased their reproductive success by a third. Both sexes modified moult timing but in different ways. Late breeding females postponed moult approximately until chick independence without compensating for delay by faster moult. Males started moult on time and overlapped it with breeding, associated with markedly slowed plumage change. Sex differences in moult score increased with lay date, but due to their respective modifications, both sexes delayed moult completion. Nonetheless, we could not detect any evidence for survival costs of late breeding. Breeding and moult of European Stonechats appear relatively flexible, despite migratory schedules and photoperiodic programs for seasonal timing. Individuals can modify seasonal behaviour in late summer, presumably depending on their condition, and may profit considerably from extended breeding.     

The effects of long‐distance migration on the evolution of moult strategies in Western‐Palearctic passerines

Although feathers are the unifying characteristic of all birds, our understanding of the causes, mechanisms, patterns and consequences of the feather moult process lags behind that of other major avian life‐history phenomena such as reproduction and long‐distance migration. Migration, which evolved in many species of the temperate and arctic zones, requires high energy expenditure to endure long‐distance journeys. About a third of Western‐Palearctic passerines perform long‐distance migrations of thousands of kilometres each year using various morphological, physiological, biomechanical, behavioural and life‐history adaptations. The need to include the largely non‐overlapping breeding, long‐distance migration and feather moult processes within the annual cycle imposes a substantial constraint on the time over which the moult process can take place. Here, we review four feather‐moult‐related adaptations which, likely due to time constraints, evolved among long‐distance Western‐Palearctic migrants: (i) increased moult speed; (ii) increased overlap between moult and breeding or migration; (iii) decreased extent of plumage moult; and (iv) moult of part or all of the plumage during the over‐wintering period in the tropics rather than in the breeding areas. We suggest that long‐distance migration shaped the evolution of moult strategies and increased the diversity of these strategies among migratory passerines. In contrast to this variation, all resident passerines in the Western Palearctic moult immediately after breeding by renewing the entire plumage of adults and in some species also juveniles, while in other species juvenile moult is partial. We identify important gaps in our current understanding of the moult process that should be addressed in the future. Notably, previous studies suggested that the ancestral moult strategy is a post‐breeding summer moult in the Western Palearctic breeding areas and that moult during the winter evolved due to the scheduling of long‐distance migration immediately after breeding. We offer an alternative hypothesis based on the notion of southern ancestry, proposing that the ancestral moult strategy was a complete moult during the ‘northern winter’ in the Afro‐tropical region in these species, for both adults and juveniles. An important aspect of the observed variation in moult strategies relates to their control mechanisms and we suggest that there is insufficient knowledge regarding the physiological mechanisms that are involved, and whether they are genetically fixed or shaped by environmental factors. Finally, research effort is needed on how global climate changes may influence avian annual routines by altering the scheduling of major processes such as long‐distance migration and feather moult.     

Non‐breeding Cackling,Ross's and Snow Geese on Baffin Island show no loss of body mass during wing moult

Non‐breeding Cackling Branta hutchinsii, Ross's Anser rossii and Lesser Snow Geese Anser caerulescens caerulescens captured during remigial moult on Baffin Island in 2015 showed no loss of body mass with moult stage, and individual variation in mass was largely explained by sex and measures of body size (tarsus length). Exceptional conditions in 2015 resulted in almost no reproductive effort or success in that year, so captured geese of all three species were likely to have been non‐breeding individuals that initiated moult early, whereas there were almost no failed or successful breeders, which would normally moult later. This suggests that in a non‐breeding year (i.e. in the absence of competition from large numbers of goslings), locally moulting geese can obtain sufficient exogenous energy to meet their needs during the flightless wing moult period without losing body mass. This also is consistent with the hypothesis that in other species of geese, accumulation of fat stores prior to, and depletion of such stores during, wing moult is adaptive and likely to be a feature of individual plasticity to meet particular needs, such as undertaking moult migration to remote sites where precise foraging and predation conditions cannot be anticipated, or where competition from more dominant individuals may restrict their access to a reliable food supply.     

Moult in the Loggerhead Shrike Lanius ludovicianus is influenced by sex,latitude and migration

We investigated moult strategies in Loggerhead Shrikes by examining first prebasic or preformative moult patterns and by assessing the general location where individual feathers were grown using stable hydrogen isotope (δ²H) analysis. We tested the relative importance of factors known to impact moult timing and pattern, including age, sex, body size, food availability and migration. Migratory Shrikes showed evidence of suspended moult, in which feathers are moulted on both the breeding and the non‐breeding grounds with a suspension of moult during migration. Extent of moult was best explained by sex, longitude, migratory behaviour and breeding‐ground latitude. Male Hatch Year (HY) Shrikes replaced more feathers on the breeding grounds prior to migration than did HY females and moulted more extensively on the breeding grounds than did females. Non‐migratory HY Shrikes underwent a more extensive preformative moult than migratory HY Shrikes. Individuals in more southerly migratory populations moulted more extensively on the breeding grounds than did those breeding further north. Our data also indicate that individuals in the northeastern populations moulted more extensively on the breeding grounds than did those in the north and southwest. Our study underlines the complex structure and variation in moult possible within species, revealing surprising levels of differentiation between sexes and age cohorts, linked to environmental factors on the breeding grounds. Our study highlights the utility of an intrinsic marker, specifically δ²H analysis, to test hypotheses regarding the evolutionary and ecological forces driving moult. Although the methodology has not commonly been applied to this area of research, our results indicate that it can provide unprecedented insight into inter‐ and intra‐specific adaptive response to constraints, whereby individuals maximize fitness.     

SOME INTERESTING RHODESIAN RECORDS—IV

Austin, G. T. 1979. Pattern and timing of moult in penduline tits (Anthoscopus). Ostrich 49:168-173.

Moult was examined in species of Anthoscopus. Second and subsequent prebasic moults were complete. Primary and rectrix moult was typical of passerines, but secondary moult was some what irregular. Moult was largely non-overlapping with breeding, although some body moult was noted during the breeding season. In southern Africa there was some regional variation in timing of moult. First year birds moulted after adults had largely completed feather replacement. This first prebasic moult was incomplete.     

Flexibility in the timing of post-nuptial moult among Red-billed Queleas Quelea quelea in Botswana in relation to the timing of breeding

The timing of primary moult of adult Red-billed Queleas Quelea quelea, captured as they were completing an unusually late breeding attempt at Francistown, northern Botswana, in June 2004, was compared with the timing of moult of birds breeding earlier in the season in north-west Botswana during two earlier years, 1971 and 1972. Differences between years in the dates when local colonies finished breeding (mid-March to late June) and between two localities in the same year (mid-March and late May) were matched by corresponding differences in the estimated dates of moult onset, ranging from mid-April to mid-June. Flexibility in the timing of moult among Red-billed Queleas in southern Africa evidently enables birds to take advantage of unusually late breeding opportunities by delaying moult onset and overlapping moult and breeding at the end of the nesting cycle. Such flexibility may also include moult interruption to permit late breeding, although its incidence in southern Africa is apparently low.     

THE RELATIONSHIP BETWEEN CLIMATE AND ANNUAL CYCLES IN THE COTINGIDAE

This survey is based primarily or the state of moult of over 4000 specimens of cotingas from all parts of the neotropical region. The seasonality of moult thus revealed is combined with existing knowledge of breeding seasons and seasonal environmental changes in an attempt to work out the broad pattern of annual cycles and their relation to climate. Within any local population the date of onset of moult may vary according to sex and age. In genera in which both sexes participate in nesting, males and females begin to moult at about the same time, or the males slightly in advance of the females. In genera with marked sexual dimorphism, in which only the female attends the nest, males may begin to moult well before females, at about the time that the latter begin egg-laying. The former group includes the genera Pachyramphus and Tityra, comprising species that are largely insectivorous, and the latter group includes the more specialized frugivorous genera. In all areas with well-marked seasonality, the ‘frugivorous group’ moults on average before the ‘Pachyramphus group’. It appears to be a general rule for first-year birds to moult earlier than older birds. A regional survey embracing all parts of the neotropical region shows that the peak of onset of moult occurs towards the end of the dry season (frugivorous group) or early in the wet season (Pachyramphus group). The changing moult seasons, strikingly in conformity with the geographical changes in the period of heaviest rainfall, are traced along a number of transects from Mexico in the north to Paraguay and Bolivia in the south. Such evidence as there is suggests that the main period of onset of moult in the frugivorous and Pachyramphus groups coincides with the period when their food is approaching or at its seasonal peak of abundance. It seems that both breeding and moult, which are almost entirely mutually exclusive, are as far as possible timed to coincide with this most favourable period; but whereas the moult takes a more or less fixed length of time the period when breeding is possible varies greatly in different species. Widely different patterns of annual cycle may result from the interaction of the two processes. Examples are given both from the cotingas and from species of other families with similar ecology. The proximate factors controlling the timing of the moult are briefly considered. It is suggested that increasing food availability is the main environmental controlling factor, and that an endogenous circannual cycle of moult must also be involved.     

BIOLOGY OF THE BANK CORMORANT,PART 2: MORPHOMETRICS,PLUMAGE, BARE PARTS AND MOULT

Cooper J. 1985. Biology of the Bank Cormorant, Part 2: Morphometrics, plumage, bare parts and moult. Ostrich 56: 79–85.

The Bank cormorant Phalacrocorax neglectus is a medium-sized, heavily built but relatively short-tailed cormorant. Males average 2 107 g, females 1794 g. There are significant differences, but overlaps, in mass, wing length, bill, tarsus and tail length between the sexes of breeding adults. The species is all black; adults have a white rump patch and white filoplumes on the head and neck in prenuptial plumage. The white rump patch is lost during breeding. Bare parts are black except for the iris (the only useful aging character) which is dark brown in juveniles, green in subadults and horizontally divided orange-brown above, green below, in adults. Partial albinism is common but leucism is not. Adults can moult while breeding but it seems likely that most moult occurs outside the breeding season. Adults and a single subadult examined exhibited Staffelmauser or stepwise moult of the primaries.     

3. THE GENERA TURDUS,GEOKICHLA, PSOPHOCICHLA,MONTICOLA AND COSSYPHA IN NYASALAND

Summary

Hanmer, D. B. 1981. Mensural and moult data of nine species of sunbird from Moçambique and Mala?i. Ostrich 52: 156–178.

Wing length, culmen length, weight and moult data are given for the Coppery Sunbird Nectarinia cuprea, Purplebanded Sunbird N. bifnsclata, Yellowbellied Sunbird N. venusta, Whitebellied Sunbird N. talatala. Grey Sunbird N. veroxii, Black Sunbird N. amethystina. Scarletchested Sunbird N. senegalensis. Collared Sunbird Anthreptes collaris and Violet-backed Sunbird A. longuemarei from Mopeia (Moçambique) and Nchalo (Mala?i). Moulting seasons, immature age at moult, the duration of primary moult, its relation to the breeding season, weight changes with moult, the breeding season, altitude and latitude, and the sequence and timing of moult in remiges and rectrices are discussed for all species except N. veroxii, N. amethystina and A. longuemarei. Softpart and plumage colour changes with age are discussed and some data are given on habits and migration.     

“THE BIRDS OF SOUTH AFRICA”

Jones, P. J. 1980. The timing of wing moult in the Greyhooded Kingfisher in Nigeria. Ostrich 51:99-106.

The post-nuptial and post-juvenile moults of the Greyhooded Kingfisher Halcyon leucocephala took place in northern Nigeria between May and November (the rainy season) after migration from the southern breeding areas. Moult of individual birds lasted between 92 and 176 days, those starting moult latest (mostly juveniles) moulting fastest. This variation may be related to food availability during moult; those starting early do so before the rainy season begins in the north and before insect numbers increase, whereas those moulting later do so during the full flush of rainy season insect availability. This variability appears to be adaptive in allowing the complete moult to be fitted into the period remaining between the end of the breeding season, which is variable, and the southward migration in the early dry season, whose timing is relatively fixed.     

NOTES ON THE PEARL-BREASTED SWALLOW HIRUNDO DIMIDIATA IN THE SOUTH-WESTERN CAPE

Earlé R. A. &; Herholdt, J. J. 1988. Breeding and moult of the Anteating Chat Myrmecocichla formicivora. Ostrich 59: 155–161.

The general breeding biology and moult of the Anteating Chat Myrmecocichla formicivora was studied in open grassveld over a two-year period. During the winter (July), groups were significantly smaller than during summer (December) (1,81 ± 0,50 versus 2,85 ± 1,35 birds per group). There was a large turnover of individuals in the study area but the total population stayed the same. The breeding season in the study area lasted from September to February but analysis of nest record cards from a larger area gave a breeding season of August-April. Two types of nests were used: 90,6% were burrows in sand banks or other excavations, but 9,4%were in the mud pellet nests of Greater Striped Swallows Hirundo cucullata (n = 53). Consecutive breeding attempts were never made in the same burrow. Clutches consisted of three, four or five eggs ([Xbar] = 3,73 ± 0,67). Incubation lasted 14–14,5-15 days. The nestling period lasted 15–18 days. Fledgling/egg breeding success was 41,8% with 48,2% of all eggs not reaching the hatching stage. Juveniles showed an unequal sex ratio of 0,57 ♂ ♂: 1,0 ♀ ♀ but adults had a nearly equal ratio (0,9 ♂ ♂: 1,0 ♀ ♀). There was a significant positive correlation between the primary moult score and the week of the seven months in which moult was recorded. Juveniles underwent a complete body moult and partial primary moult 3–4 months after fledging.     

MEASUREMENTS AND MOULT OF FIVE SPECIES OF BULBUL FROM MOÇAMBIQUE AND MALAŴI

Summary

Hanmer, D. B., 1978. Measurements and moult of five species of bulbul from Moçambique and Mala?i. Ostrich 49:116-131.

The wing length, weight and moult of five species of bulbul, Blackeyed Pycnonotus barbatus, Sombre Andropadus importunus, Yellowbellied A. flaviventris, Terrestrial Phyllas-trephus terrestris and Yellowspotted Nicator Nicator gularis, are given for two localities in tropical lowland (Mopcia, Moçambique and Nchalo, Mala?i). The characters identifying immatures and the length of time these are retained, are given with reference to skull pneumatization, retrapped birds and the breeding season, for Pycnonotus barbatus, Andropadus intportunus and Phyllastrephus terrestris. Weights are compared with some published for other parts of Africa. The months during which moult occurred are given. Duration and timing of primary moult and its relation to the breeding season, are given for Pycnonotus barbatus, Andropadus importunus and Phyllastrephus terrestris. The age at which immatures moult is given for these three species. Instances of interrupted moult are mentioned.     

Breeding barnacle geese in Kolokolkova Bay,Russia: number of breeding pairs,reproductive success and morphology

We report the results of an expedition to a barnacle-goose (Branta leucopsis) breeding area in Kolokolkova Bay, west of the lower Pechora delta in northern Russia, undertaken in July 2002. In total, 6 breeding colonies were found within the study area, harbouring 1,324 nests. Mean clutch size was 2.77±0.10 but may have been underestimated because of nest predation. Nest predation was high and correlated with the density of breeding gulls, Larus. The 2002 season was relatively cold and peak hatch occurred late, on 14 July. More than 11,000 barnacle geese were found to moult in the area which, together with the large number of nests found, emphasises the importance of Kolokolkova Bay for barnacle geese. Adult barnacle geese (341) were captured, marked and measured during their annual wing moult. Birds with broods started to moult approximately 2 weeks later than non- and failed breeders. Weight loss during moult was 3 times as rapid as reported for barnacle geese breeding in the Baltic, and a large cost of reproduction seemed to exist in the form of reduced body weight at the onset of moult for birds leading broods. Work in the area will continue over the coming years to document and explain the differences in major life-history parameters, dynamics and environmental effects between arctic and temperate breeding barnacle-goose populations.     

Identifying the winter grounds of the recently described Barbary Reed Warbler (Acrocephalus baeticatus ambiguus)

The Iberian and North African populations of reed warblers have been described recently as a separate taxon, ambiguus, forming a sister clade to the Sahelian subspecies minor of African Reed Warbler Acrocephalus baeticatus. Although the breeding range of ambiguus has been identified, the migration strategy of its populations remained unknown. We deployed geolocators and sampled the innermost primary from breeding adults in Spain for stable hydrogen (δ²H) analyses and also analysed stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes in feathers collected in two reed warbler taxa (Acrocephalus scirpaceus and Acrocephalus baeticatus ambiguus) in Morocco, to identify the moulting and wintering sites of these populations. Ring recoveries, geolocator tracks and probabilistic assignments to origin from δ²H values indicate that Spanish ambiguus are likely to moult south of the Sahara and winter in West Africa, probably from Mauretania to southern Mali and Ivory Coast. Moroccan ambiguus, however, undergo post-breeding moult north of the Sahara, and possibly then migrate to West Africa. With other populations of ambiguus breeding from Algeria to Libya and probably wintering further east in the Sahelian belt, the Barbary Reed Warbler can therefore be considered a trans-Saharan migrant, with a post-breeding moult strategy that varies between populations, and probably structured according to breeding latitude.     

Timing of breeding and primary moult of the Masked Weaver Ploceus velatus in the summer and winter rainfall regions of South Africa

Oschadleus, H.D., Underhill, G.D. & Underhill, L.G. 2000. Timing of breeding and primary moult of the Masked Weaver Ploceus velatus in the summer and winter rainfall regions of South Africa. Ostrich 71 (1 & 2): 91–94.

Timing of breeding and moult is analysed in the Masked Weaver Ploceus velatus. It is common throughout southern Africa, which is largely a summer rainfall area. This species expanded its range into the Western Cape, a winter rainfall region, in the twentieth century. The peak breeding period is one month earlier in the winter rainfall area (September to November) than in the summer rainfall area (October to December). The mean starting date of primary moult is one month earlier in the winter rainfall area (9 January) than in the summer rainfall area (15 February). The duration of primary moult is similar in both regions (74 days in the winter rainfall area and 80 days in the summer rainfall area).     

PRIMARY MOULT,WEIGHT AND BREEDING CYCLES OF THE ROCK PIGEON ON DASSEN ISLAND

Cooper, J. 1975. Primary moult, weight and breeding cycles of the Rock Pigeon on Dassen Island. Ostrich 46:154-156.

The primary moult season of the adult Rock Pigeon Columba guinea on Dassen Island is spread over at least nine months. Individual duration is estimated at eight months. Adult birds were heaviest in the winter months outside the breeding season. Overlap between the breeding and moulting seasons occurred and evidence was obtained of incubating birds with active primary moult. Juveniles were lighter than adults. Adults fed on the mainland and probably made daily flights there.     

Greater energy stores enable flightless moulting geese to increase resting behaviour

Many species of waterfowl undergo a post‐breeding simultaneous flight feather moult (wing moult) which renders them flightless and vulnerable to predation for up to 4 weeks. Here we present an analysis of the correlations between individual time‐budgets and body mass states in 13 captive Barnacle Geese Branta leucopsis throughout an entire wing moult. The daily percentage of time spent resting was positively correlated with initial body mass at the start of wing moult. Behaviour of individual birds during wing moult is dependent on initial physiological state, which may in turn be dependent on foraging ability; the storage of energy before the start of wing moult will help birds to reduce exposure to the dangers of predation.     

The timing of gonadal development and moult in three raptors with different breeding seasons: effects of gender, age and body condition

Differences between species in breeding seasons are thought to be mediated through differences in their reproductive physiology. Little is known about how the timing and duration of gonadal maturation varies between raptor species, how the timing of moult relates to the gonadal cycle, whether the timing and degree of sexual maturation varies between juveniles and adults or whether body condition has a significant effect. To address these questions, data on gonadal size and moult for adults and juveniles of both sexes of three raptor species were extracted from the Predatory Bird Monitoring Scheme (based on birds found dead by members of the public). The three species, Sparrowhawk Accipiter nisus, Kestrel Falco tinnunculus and Barn Owl Tyto alba, have different ecologies – diurnal bird predator, diurnal mammal predator and nocturnal mammal predator, respectively. All are single‐brooded but have different breeding seasons. The duration of gonadal maturation was markedly different between the species. Barn Owls showed the earliest maturation and the latest gonad regression, and Sparrowhawks the latest maturation and earliest gonad regression. Kestrels were intermediate. In males of all species, the testes remained fully mature throughout their respective breeding seasons. In females, the ovaries remained partially mature throughout the breeding season. Moult started slightly earlier in Sparrowhawks than in Kestrels and coincided with gonadal regression in the two species. Although females of the two species started to moult earlier than males, moult duration was similar between the sexes. Barn Owls showed no distinct annual pattern of moult. In juveniles of all three species, the gonads were smaller than in adults throughout spring and started to mature later. Gonad size in birds that had starved tended to be smaller than in birds dying from other causes, but did not influence the difference in gonad mass between adults and juveniles and between seasons. Body condition had no effect on moult. Whilst ecology has led to the evolution of different breeding seasons, differences between species, and between adults and juveniles, are mediated through adaptive differences in their reproductive physiology.     

Skuas (Stercorarius spp.) moult body feathers during both the breeding and inter‐breeding periods: implications for stable isotope investigations in seabirds

Seabirds are mostly thought to moult during the inter‐breeding period and the isotopic values of their feathers are often therefore assumed to relate to their assimilated diet during such periods. We observed Brown Skuas Stercorarius antarcticus lonnbergi and South Polar Skuas Stercorarius maccormicki moulting on a breeding site at King George Island, Antarctica. This raises concerns about the reliability of using stable isotopes in feathers to infer feeding localities of birds during the inter‐breeding period. We analysed the δ¹³C and δ¹⁵N values of growing and fully grown body feathers collected from the same individuals. For both species, δ¹³C values of growing feathers indicated feeding areas in the Antarctic zone (breeding grounds), whereas most fully grown feathers (100% for South Polar Skuas and 93.3% for Brown Skuas) could be assigned to northern latitudes (non‐breeding grounds). However, a few fully grown body feathers of Brown Skuas (6.7% of the feathers, belonging to two birds) showed isotopic values that indicated moult in the Antarctic zone. As the growth period of those feathers was unknown, they could not be used with confidence to depict the foraging behaviour of the birds during the non‐breeding period. Although precautions must be taken when inferring dietary information from feathers in seabirds where the moulting pattern is unknown, this study shows that if the development stage of a feather (growing/fully grown) is identified, then dietary information from both breeding and non‐breeding seasons can be obtained on the same individual birds.     

Adjustment of pre‐moult foraging strategies in Macaroni Penguins Eudyptes chrysolophus according to locality,sex and breeding status

The annual moult creates the highest physiological stress during a penguin's breeding‐cycle and is preceded by a period of hyperphagia at sea. Although crucial to individual survival, foraging strategies before moult have been little investigated in keystone marine consumers in the Southern Ocean. The Macaroni Penguin Eudyptes chrysolophus demonstrates how individuals may adjust their foraging strategies during this period in line with constraints such as potential intraspecific competition between localities, foraging ability between dimorphic sexes and timing at sea between breeding and non‐breeding population components. We recorded pre‐moult behaviour at sea for 22 Macaroni Penguins from Crozet and Kerguelen Islands (southern Indian Ocean) during 2009 and 2011, using light‐based geolocation and stable isotope analysis. Penguins were distributed in population‐specific oceanic areas with similar surface temperatures (3.5 °C) south of the archipelagos, where they foraged at comparable trophic levels based on stable isotopes of their blood. Bayesian ‘broken stick’ modelling with concurrent analysis of seawater temperature records from the animal‐borne devices showed that within each population, females remained 6 days longer than males in the colder waters before heading back towards their colonies. Finally, 17 other non‐breeding individuals that moulted earlier had a higher mean blood δ¹⁵N value than did post‐breeding birds, meaning that early moulters probably fed more on fish than did late moulters. Our findings of such adjustments in foraging strategies developed across locality, sex and breeding status help understanding of the species' contrasted pre‐moult biology across its range and its ecology in the non‐breeding period.