Variability of a consistent trait: The size of the white wing patch in European Stonechats (Saxicola rubicola rubicola)

Numerous bird species exhibit striking white patches on their plumages that have been reported as signals of individual quality in the context of sexual selection. Whereas differences in white plumage traits between individuals have been well studied, phenotypic variation and the factors affecting their consistency within the individual have received less attention. Here, we studied the consistency in the size of the white wing patch and its components of variation in a Spanish population of European Stonechats Saxicola rubicola rubicola monitored over 6 years (2007–2012). The wing patch was larger in males than in females for all age-classes, and first-year individuals exhibited smaller wing patches than older birds, particularly males. This trait also varied within the plumage-year at both population and individual level, with slight changes from the moulting period to the breeding season and a sharp decrease afterwards. In addition, patch size varied both between and within individuals across years. The daily mean temperature experienced in the month immediately prior to the moult (i.e. May) had a positive effect on male wing patch size. In females, however, no variable was found to explain wing patch variation satisfactorily. Despite being a dynamic trait throughout the moults, baseline and adjusted repeatabilities indicated a moderately high consistency for white wing patch size in both sexes. Therefore, the white wing patch in Stonechats is a dynamic trait that changes throughout individuals' lives, but is also a consistent trait at the individual level. These results indicate that this depigmented patch could convey information about both the long-term quality and the current state of individuals, harmonizing the existence of intra-individual changes in the size of sexual traits with its potential role as a genetic quality indicator.     

Optimal moult strategies in migratory birds

Avian migration, which involves billions of birds flying vast distances, is known to influence all aspects of avian life. Here we investigate how birds fit moult into an annual cycle determined by the need to migrate. Large variation exists in moulting patterns in relation to migration: for instance, moult can occur after breeding in the summer or after arrival in the wintering quarters. Here we use an optimal annual routine model to investigate why this variation exists. The modelled bird's decisions depend on the time of year, its energy reserves, breeding status, experience, flight feather quality and location. Our results suggest that the temporal and spatial variations in food are an important influence on a migratory bird's annual cycle. Summer moult occurs when food has a high peak on the breeding site in the summer, but it is less seasonal elsewhere. Winter moult occurs if there is a short period of high food availability in summer and a strong winter peak at different locations (i.e. the food is very seasonal but in opposite phase on these areas). This finding might explain why only long-distance migrants have a winter moult.     

Prolonged and flexible primary moult overlaps extensively with breeding in beach‐nesting Hooded Plovers Thinornis rubricollis

We present the first report of complete overlap of breeding and moult in a shorebird. In southeastern Australia, Hooded Plovers Thinornis rubricollis spend their entire lives on oceanic beaches, where they exhibit biparental care. Population moult encompassed the 6‐month breeding season. Moult timing was estimated using the Underhill–Zucchini method for Type 2 data with a power transformation to accommodate sexual differences in rates of moult progression in the early and late stages of moult. Average moult durations were long in females (170.3 ± 14.2 days), and even longer in males (210.3 ± 13.5 days). Breeding status was known for most birds in our samples, and many active breeders (especially males) were also growing primaries. Females delayed the onset of primary moult but were able to increase the speed of moult and continue breeding, completing moult at about the same time as males. The mechanism by which this was achieved appeared to be flexibility in moult sequence. All moult formulae fell on one of two linked moult sequences, one faster than the other. The slower sequence had fewer feathers growing concurrently and also had formulae indicating suspended moults. Switching between sequences via common formulae is possible at many points during the moult cycle, and three of 12 recaptures were confirmed to have switched sequences in the same moult season. Hooded Plovers thus have a prolonged primary moult with the flexibility to change their rate of moult; this may facilitate high levels of replacement clutches that are associated with passive nest defence and low reproductive success.     

Uropygial gland size,feather holes and moult performance in the House Sparrow Passer domesticus

Feather holes represent damage to the plumage of birds and are correlated with delayed moult. Uropygial gland size is negatively correlated with feather holes. Consequently, it was predicted that birds with smaller uropygial glands would have more feather holes, and that this would affect moult performance. I examined this prediction in the House Sparrow Passer domesticus. Individuals with smaller uropygial glands had more feather holes, and those with more feather holes moulted later and faster. Therefore, uropygial gland size seemed to affect moult performance via its effect on feather holes. Uropygial gland size may have a positive effect on plumage quality, through a negative effect on feather holes, and therefore on moult timing and speed.     

Factors influencing the evolution of moult in the non‐breeding season: insights from the family Motacillidae

The number of moults per annual cycle and their final spatial pattern (i.e. topography) show high interspecific variation in the order Passeriformes. Factors behind this variability remain obscure, especially for variability in spatial pattern among species. Here, we explored the relative influence of ten ecological, ontogenetic, social and sexual factors on the evolution of autumn moult (feather replacement largely undertaken by migratory species, which is not necessarily an independent episode within their moult cycle) and prealternate moult among Northern Hemisphere species of the family Motacillidae using phylogenetically controlled analyses, ancestral state reconstruction and analyses of correlated evolution. The results strongly support the presence of prealternate moult and absence of autumn moult as ancestral states in this family. A high rate of change between related species indicates phylogenetic independence among prealternate moult patterns and examined factors. Migration distance and gregariousness are the most important factors influencing prealternate moult evolution, and point toward natural selection and sociality as the most important evolutionary drivers of prealternate moult in Motacillidae. Breeding latitude, seasonal plumage change, winter plumage conspicuousness, sexual dichromatism, plumage maturation and extent of preformative moult show a minor influence, and suggest that ontogeny and sexual selection may have played a limited role in shaping prealternate moult in Motacillidae.     

Frequency of complete moult in adult and juvenile Great Reed Warblers(Acrocephalus arundinaceus) in Spain

Summary We examined a large data set of adult and juvenile Great Reed Warblers (Acrocephalus arundinaceus), caught over five years, to study the occurrence of complete moult in a Spanish population. A total of 27 adults and 5 juveniles were in active moult. The proportion of birds moulting each year varied between 0.03 and 0.19. Nearly half the adults were finishing moult. At least one bird was certainly of local origin, rather than on passage, and four more were moulting when retrapped in subsequent years, suggesting that they too were resident breeders in the area. Some other birds were probably resident also, because they were in moult when caught at the beginning of August. Two of the five juveniles finished their moult in the area. In conclusion, our data seem to show that complete moult before departure to the wintering quarters is a regular process undertaken by some of the breeding population of Great Reed Warblers in north-east Spain.

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Das Vorkommen von Vollmauser bei Alt- und Jungvögeln des Drosselrohrsängers(Acrocephalus arundinaceus) in Spanien

Zusammenfassung Von 837 zwischen 1992 und 1996 im Ebro Delta gefangenen Drosselrohrsängern mauserten 27 Alt-und 5 Jungvögel. Der jährlichen Anteil mausernder Vögeln betrug zwischen 0,03 und 0,19%. Etwa die Hälfte der mausernden Altvögel und zwei der Jungvögel beendeten ihre Mauser im Untersuchungsgebiet. Diese Daten zeigen, daß Drosselrohrsänger in Spanien wenigstens teilweise auch schon vor Beginn des Wegzuges eine Vollmauser durchführen.

     

Contrasting strategies for wing‐moult and pre‐migratory fuelling in western and eastern populations of Common Whitethroat Sylvia communis

Trade‐offs between moult and fuelling in migrant birds vary with migration distance and the environmental conditions they encounter. We compared wing moult and fuelling at the northern and southern ends of migration in two populations of adult Common Whitethroats Sylvia communis. The western population moults most remiges at the breeding grounds in Europe (e.g. Poland) and migrates 4000–5000 km to western Africa (e.g. Nigeria). The eastern population moults all remiges at the non‐breeding grounds and migrates 7000–10 000 km from western Asia (e.g. southwestern Siberia) to eastern and southern Africa. We tested the hypotheses that: (1) Whitethroats moult their wing feathers slowly in South Africa, where they face fewer time constraints than in Poland, and (2) fuelling is slower when it coincides with moulting (Poland, South Africa) than when it occurs alone (Siberia, Nigeria). We estimated moult timing of primaries, secondaries and tertials from moult records of Polish and South African Whitethroats ringed in 1987–2017 and determined fuelling patterns from the body mass of Whitethroats ringed in all four regions. The western population moulted wing feathers in Poland over 55 days (2 July–26 August) at a varying rate, up to 13 feathers simultaneously, but fuelled slowly until departure in August–mid‐September. In Nigeria, during the drier period of mid‐February–March they fuelled slowly, but the fuelling rate increased three‐fold in April–May after the rains before mid‐April–May departure. The eastern population did not moult in Siberia but fuelled three times faster before mid‐July–early August departure than did the western birds moulting in Poland. In South Africa, the Whitethroats moulted over 57 days (2 January–28 February) at a constant rate of up to nine feathers simultaneously and fuelled slowly from mid‐December until mid‐April–May departure. These results suggest the two populations use contrasting strategies to capitalize on food supplies before departure from breeding and non‐breeding grounds.     

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.     

Age‐specific variation in relationship between moult and pre‐migratory fuelling in Wood Sandpipers Tringa glareola in southern Africa

Trans‐equatorial avian migrants tend to breed, moult and migrate – the main energy‐requiring events in their lifecycle – at different times. Little is known about the relationship between wing moult and pre‐migratory fuelling in waders on their non‐breeding grounds, where time is less constrained than during their brief high‐latitude breeding season. We determined age‐related strategies of Wood Sandpipers Tringa glareola to balance the energetic demands of primary moult against pre‐migratory fuelling in southern Africa by analysing body mass and primary moult at first capture of 1721 birds mist‐netted in 1972–96 at waterbodies in Zimbabwe. Adults moulted all their primaries in August–December, but immatures underwent a supplemental moult of varying numbers of outer primaries in December–April, close to departure. We used locally weighted linear regression to estimate trends in Wood Sandpiper body mass from 1 July to 1 May. They maintained low mass from arrival in July–September to February–early March. Adults fuelled from 10 February to 1 May at a mean rate of 0.25 g/day (sd = 0.16). Most adults (98%) began fuelling 10–75 days after completing primary moult. Immatures fuelled from 4 March to 13 April at 0.24 g/day (sd = 0.14). They used varying strategies depending on their condition: a brief gap between moult and fuelling; an overlap of these processes near departure, leading to slower fuelling; or skipping fuelling altogether and staying in southern Africa for a ‘gap year’. Immatures moulting three or five outer primaries fuelled more slowly than post‐moult birds. Immatures moulting four outer primaries started fuelling 3 weeks later but at a higher rate than did post‐moult birds of this group. In post‐moult immatures, the later they ended moult, the later and faster they fuelled. The heaviest adults and immatures using all moult patterns accumulated fuel loads of c. 50% of lean body mass, and could potentially cross 2397–4490 km to reach the Great Rift Valley in one non‐stop flight. Immatures were more flexible in the timing and extent of moult and in the timing and rate of fuelling than adults. This flexibility enables inexperienced Wood Sandpipers to cope with inter‐annual differences in feeding conditions at Africa's ephemeral inland waterbodies.     

Flexibility in the timing of post‐breeding moult in passerines in the UK

Higher temperatures resulting from climate change have led to predictions that the duration of the breeding season of many temperate bird species may be changing. However, the extent to which breeding seasons can be altered will also depend on the degree of flexibility in processes occurring at other points in the annual cycle. In particular, plasticity in the timing of post‐breeding moult (PBM) could facilitate changes in the timing of key events throughout the annual cycle, but little is known about the level of within‐ and between‐species plasticity in PBM. As part of the British Trust for Ornithology (BTO) Ringing Scheme, many ringers routinely record moult scores of flight feathers, and these can be used to provide information on the annual progression of PBM for a range of species. Here we use ringing data to investigate patterns of PBM in 15 passerines, as well as data from the BTO Nest Record Scheme to relate these differences to the timing of breeding of these species across the UK. We find considerable variation in both the mean start (19 May–29 July) and duration (66–111 days) of PBM between species, but find no evidence that species starting PBM later in the season complete it any faster. However, there is considerable within‐species variation in PBM, particularly for multi‐brooded species; PBM starts later and is completed in less time when the duration of the breeding season (difference between first and last nests) is longer. This implies that a later end to breeding can be compensated for by faster PBM, and that advances in breeding could lead to earlier and slower PBM. Our findings suggest that adaptation of PBM in response to climate‐mediated changes in the timing and duration of the breeding season is possible. However, the requirement to complete PBM prior to migration or the onset of winter might constrain the extent to which breeding seasons can lengthen, especially for later nesting species.     

Testosterone and corticosterone during the breeding cycle of equatorial and European stonechats (Saxicola torquata axillaris and S. t. rubicola)

Northern-temperate male birds show seasonal changes in testosterone concentrations with a peak during the breeding season. Many tropical birds express much lower concentrations of testosterone with slight elevations during breeding. Here we describe testosterone and corticosterone concentrations of male stonechats from equatorial Kenya during different substages of breeding and molt. This tropical species has a short breeding season of approximately 3 months. We compare their hormone concentrations to previously published data of males of a northern-temperate relative, the European stonechat, also a seasonal breeder but with a breeding season of approximately 5 months. Equatorial stonechats show a pronounced peak of testosterone during the nest-building and laying stage. During all other stages, testosterone concentrations are low, similar to other year-round territorial tropical bird species. Corticosterone concentrations peak also during the nest-building and laying stage suggesting that this period of maximum female fecundity is a demanding period for the male. Equatorial stonechats have significantly lower concentrations of testosterone than European stonechats during all stages, except during the nest-building and laying stage. During this stage of maximum female fertility, testosterone levels tend to be higher in equatorial than in European stonechats. Our results suggest that equatorial stonechats belong to a group of tropical bird species that are characterized by a short breeding season and a brief high peak of testosterone during the female's fertile period. Such brief, but substantial peaks of testosterone may be common in tropical birds, but they may easily be missed if the exact breeding stage of individual birds is not known.     

Sex‐specific patterns of fuelling and pre‐breeding body moult of Little Stints Calidris minuta in South Africa

The timing and duration of each stage of the life of a long‐distance migrant bird are constrained by time and resources. If the parental roles of males and females differ, the timing of other life stages, such as moult or pre‐migratory fuelling, may also differ between the sexes. Little is known about sexual differences for species with weak sexual dimorphism, but DNA‐sexing enables fresh insights. The Little Stint Calidris minuta is a monomorphic long‐distance migrant wader breeding in the Arctic tundra. Males compete for territories and perform elaborate aerial displays. Females produce two clutches a season. Each sex may be a bigamist and incubate one nest a season, each with a different partner. We expect that these differences in breeding behaviour entail different preparations for breeding by males and females, so we aimed to determine whether Little Stints showed any sex differences in their strategies for pre‐breeding moult and pre‐migratory fuelling at their non‐breeding grounds in South Africa. We used body moult records, wing length and body mass of 241 DNA‐sexed Little Stints that we caught and ringed between 27 January and 29 April in 2008–2018 at two neighbouring wetlands in North West Province, South Africa. For each individual we assessed the percentage of breeding plumage on its upperparts and took blood samples for DNA‐sexing. We calculated an adjusted Body Moult Index and an adjusted Wing Coverts Moult Index, then used the Underhill–Zucchini moult model to estimate the start dates and the rate of body moult in males and females. We estimated the changes in the sex ratio of the local population during their stay in South Africa, and also estimated the timing and rate of pre‐migratory fuelling and the potential flight ranges for males and females. The males started body moult on average on 7 February and the females on 12 February, but the sexes did not differ in their timing of wing covert moult, which started on average on 10 February. In January to mid‐February, males constituted c. 57% of the population, but their proportion declined afterwards, indicating an earlier departure than females. We estimated that both sexes began pre‐migratory fuelling on average on 15 March. The sexes did not differ in fuelling rate, but most females stayed at the non‐breeding site longer than the males, and thus accumulated more fuel and had longer potential flight ranges. These patterns of moult and fuelling suggest sex differences in preparations for breeding. We suggest that the males depart from South Africa earlier but with smaller fuel loads than the females to establish breeding territories before the females arrive. We conclude that for each sex the observed trade‐offs between fuelling and moult at the non‐breeding grounds are precursors to different migration strategies, which in turn are adaptations for their different roles in reproductive behaviour.     

Seasonally divided moult in the Barred Warbler (Sylvia nisoria) is an endogenously controlled strategy

The seasonal timing of moult in migratory birds is an adaptation to cope with time constraints in the annual cycle. Kiat and Izhaki analysed moult patterns in Palaearctic passerines and rejected the proposition that seasonally divided moult is an endogenously controlled strategy. Instead, they advocated the view that it occurs due to a flexible and opportunistic timing of moult. In contrast, we argue that Kiat and Izhaki’s analysis is flawed and that they overlooked several important facts about moult in the Barred Warbler Sylvia nisoria and other species showing seasonally divided moult. These include the facts that juveniles replace a few secondaries already in their first winter, and that the moult sequences of primaries and secondaries are decoupled compared with the typical passerine complete moult sequence. We argue that seasonally divided moult is an adaptive strategy that is largely under endogenous control.     

Independent trophic behavior and breeding success of a resident flycatcher and a coexisting migratory congener

Historical biases towards the study of Palearctic and Nearctic bird migration systems has resulted in a dearth of information on the ecology of intratropical migrants, which likely also play important ecological roles within their communities. For instance, there is little information on the foraging ecology and breeding biology of co‐occurring migrants and residents within the intratropical migratory system. Thus, we used two congeneric flycatchers – the resident Plain‐crested Elaenia Elaenia cristata (Aves: Tyrannidae), and the migrant Lesser Elaenia E. chiriquensis (Tyrannidae) – to check whether either of them showed foraging niche plasticity mediating their coexistence and the effects of such syntopy on their breeding behavior and fitness. To do so, we used an ensemble of methods that included the evaluation of food resource phenology, foraging behavior observations, stable isotope ratio analysis in birds’ tissues, and nest monitoring. Our results confirmed that residents have foraging behavior plasticity but reveals its trophic behavior and breeding success is independent of the coexistence with migrants. Since such results depart from the predictions based on the current (and often Northern‐biased) theories of bird migration, we discuss alternative hypotheses explaining our findings, including the influence of physiology on diet and the behavioral responses of nest predators. Once there is a clear knowledge gap about the interspecific interactions between residents and intratropical migrants, our study represents a simple, yet important, step towards understanding the mechanisms underlying this system.     

Ecological correlates in the evolution of moult strategies in Western Palearctic passerines

Moult is an important process in the life cycle of birds. Passerines differ widely in the number, seasonality and extension of moult episodes, but the incidence of birds ecology on this variation remains largely uninvestigated. We analysed the patterns of moult in European passerines in relation to their distribution, migration and sexual dichromatism. Longer migrations and southern wintering quarters were characteristic of species with complete moults in summer and an additional moult in winter. The main moult in species with larger seasonal changes in sexual dimorphism tended to be scheduled just before the start of the breeding season, suggesting a link between sexual selection and the timing of moult. These patterns strongly support the importance of migration and dichromatism on the evolution of moult strategies.     

Divergent rectrix moult: the implications and conditions of moult sequence

Wing and tail morphology strongly affect flight performance which may consequently decline during feather moult due to the creation of feather gaps in the flight‐surface. Hence, the size and shape of moult‐related gaps may directly affect flight capacity. Here, I examined the divergent rectrix moult sequence compared to the more common distal moult sequence. In the divergent moult, the focus of rectrix moult is shifted from the tail centre (R₁; rectrices numbered distally from mid‐tail outward) to another rectrix (R₂ or R₃), and then rectrices are moulted bidirectionally, towards the tail centre and outwards. The result of this moult sequence is the splitting of the tail gap into multiple smaller gaps. Using a large moult database including 5669 individuals of 47 Western Palaearctic passerine species, I found evidence of divergent moult sequence for only seven species. Using comparative and experimental approaches, I found that the divergent rectrix sequence is correlated with higher moult speed and lower aerodynamic cost. Furthermore, the divergent rectrix sequence is more common among adults than juveniles. This work focused on the feather moult sequence – a seldom studied aspect of the avian life‐history. I propose that moult‐related aerodynamic costs may be an important evolutionary factor not only in moult speed, but also in moult sequence.     

Climate change and fitness components of a migratory bird breeding in the Mediterranean region

The increase in spring temperatures in temperate regions over the last two decades has led to an advancing spring phenology, and most resident birds have responded to it by advancing their onset of breeding. The pied flycatcher (Ficedula hypoleuca) is a long‐distance migrant bird with a relatively late onset of breeding with respect to both resident birds and spring phenology in Europe. In the present correlational study, we show that some fitness components of pied flycatchers are suffering from climate change in two of the southernmost European breeding populations. In both montane study areas, temperature during May increased between 1980 and 2000 and an advancement of oak leafing was detected by using the normalized difference vegetation index (NDVI) to assess tree phenology. This might result in an advancement of the peak in availability of caterpillars, the main prey during the nestling stage. Over the past 18 yr, the time of egg laying and clutch size of pied flycatchers were not affected by the increase in spring temperatures in these Mediterranean populations. However, this increase seems to have an adverse effect on the reproductive output of pied flycatchers over the same period. Our data suggest that the mismatch between the timing of peak food supply and nestling demand caused by recent climate change might result in a reduction of parental energy expenditure that is reflected in a reduction of nestling growth and survival of fledged young in our study populations. The data seem to indicate that the breeding season has not shifted and it is the environment that has shifted away from the timing of the pied flycatcher breeding season. Mediterranean pied flycatchers were not able to advance their onset of breeding, probably, because they are constrained by their late arrival date and their restricted high altitude breeding habitat selection near the southern border of their range.     

Effects of post-breeding moult and energetic condition on timing of songbird migration into the tropics

Each autumn billions of songbirds migrate between the temperate zone and tropics, but little is known about how events on the breeding grounds affect migration to the tropics. Here, we use light level geolocators to track the autumn migration of wood thrushes Hylocichla mustelina and test for the first time if late moult and poor physiological condition prior to migration delays arrival on the winter territory. Late nesting thrushes postponed feather moult, and birds with less advanced moult in August were significantly farther north on 10 October while en route to the tropics. Individuals in relatively poor energetic condition in August (high β-Hydroxybutyrate, low triglyceride, narrow feather growth bars) passed into the tropics significantly later in October. However, late moult and poor pre-migratory condition did not result in late arrival on the winter territory because stopover duration was highly variable late in migration. Although carry-over effects from the winter territory to spring migration may be strong in migratory songbirds, our study suggests that high reproductive effort late in the season does not impose time constraints that delay winter territory acquisition.     

Weak breeding seasonality of a songbird in a seasonally arid tropical environment arises from individual flexibility and strongly seasonal moult

In some tropical birds, breeding seasonality is weak at the population level, even where there are predictable seasonal peaks in environmental conditions. It therefore remains unclear whether individuals are adapted to breeding at specific times of the year or flexible to variable environmental conditions. We tested whether the relative year‐round breeding activity of the Common Bulbul Pycnonotus barbatus arises due to within‐individual variability in breeding dates. We collected data from 827 birds via mist‐netting over 2 years with corresponding local weather data. We used a combination of climate envelope and generalized linear mixed models to explore how the timing of breeding is influenced by time of year, individual variation, rainfall and temperature in a West African savannah where seasonal precipitation determines annual variation in environmental conditions. We also pooled 65 breeding records from 19 individuals recorded between 2006 and 2017 based on brood patch occurrence and behavioural observation to compare within‐individual and population variability in breeding dates. We show that the breeding dates of individuals may be as variable as for the population as a whole. However, we observed a seasonal peak in juvenile occurrence that varies significantly between years. Models suggest no relationship between nesting and moult, and within‐year variation in rainfall and temperature, and birds were unlikely to breed during moult but may do so afterwards. Moult was very seasonal, correlating strongly with day length. We suggest that because environmental conditions permit year‐round breeding, and because reproductive output is subject to high predation risk, there is probably a weak selection for individuals to match breeding with variable peak conditions in the environment. Instead, moult, which always occurs annually and successfully, is probably under strong selection to match variable peak conditions in the environment so that long‐term survival ensures future reproduction.     

White tail spots in breeding Barn Swallows Hirundo rustica signal body condition during winter moult

The determinants and function of pigmentation of feathers and other tissues have been the focus of a large number of studies, particularly with respect to socio‐sexual communication. However, many birds exhibit depigmented white spots or bars on their feathers whose function is poorly understood. Here we assess whether white feather spots reflect phenotypic condition at the time of moult by investigating the covariation between spot size or shape and condition‐dependent feather growth rate, as gauged by width of the growth bars on the tail feathers of Barn Swallows. We found that feathers with higher growth rates had larger, less rounded white spots. In addition, variance in spot perimeter for a given spot area was larger in males than in females. This study is the first to provide evidence that features of white markings on feathers directly reflect body condition at the time of moult and can therefore reliably signal phenotypic quality in the context of socio‐sexual communication. In addition, the study highlights the potential communication function of the shape and not just the size of colour signals.