Moult speed affects structural feather ornaments in the blue tit

Growing evidence suggests that structural feather colours honestly reflect individual quality or body condition but, contrary to pigment‐based colours, it is not clear what mechanism links condition to reflectance in structural feather colours. We experimentally accelerated the moult speed of a group of blue tits (Cyanistes caeruleus) by exposing them to a rapidly decreasing photoperiod and compared the spectral characteristics of their structural feather colours with those of control birds. Blue tits were sexually dimorphic on the UV/blue crown and on the white cheek feathers. Moult speed, however, dramatically reduced brightness and the saturation only on the UV/blue crown feathers, whereas structural white on the cheek feathers was basically unaffected by moult speed. Given that the time available for moulting is usually confined to the period between the end of the breeding season and migration or wintering, UV/blue colours, but not structural white, may convey long‐term information about an individual’s performance during the previous breeding season. The trade‐off between fast moulting and structural colour expression may represent a previously unrecognized selective advantage for early‐breeding birds.     

Nestlings’ carotenoid feather ornament affects parental allocation strategy and reduces maternal survival

In some birds, feather ornaments are expressed in nestlings well before sexual maturation, possibly in response to parental favouritism towards high‐quality offspring. In species with synchronous hatching, in which nestling ornaments may vary more among than within broods, parents may use this information to adjust their parental allocation to the current brood accordingly. We tested this hypothesis in the rock sparrow, in which a sexually selected yellow feather ornament is also expressed in nestlings. We experimentally enlarged nestlings’ breast patch in a group of broods and sham‐manipulated another group of control broods. Nestlings with enlarged ornament were fed more frequently and defended more actively from a dummy predator than their control counterparts. Mothers from the enlarged group were more likely to lay a second clutch and showed a reduced survival to the next breeding season. These results provide one of the first evidences of differential parental allocation among different broods based directly on nestlings’ ornamentation, and the first, to our knowledge, to show a reduction in maternal survival.     

Breeding plumage honestly signals likelihood of tapeworm infestation in females of a long-distance migrating shorebird, the bar-tailed godwit

The indicator mechanism for sexual selection proposed by Hamilton and Zuk (i.e. that sexually selected ornaments signal parasite resistance) has received rather little observational support, and none in the case of long-distance migrant birds. Here we present a test by examining the association between helminth infestations and breeding plumage quality in bar-tailed godwits Limosa lapponica taymyrensis during their spring staging period in the Wadden Sea, The Netherlands. After a non-stop flight from West Africa, these shorebirds refuel in the Wadden Sea in preparation for a second flight to the central Siberian Arctic breeding grounds. Earlier studies have shown that only relatively heavy and well ornamented birds carry out a "top-up" moult during stopover, in which part of the contour feathers recently grown in West Africa are replaced by even fresher ones. Active body moult was therefore taken as the primary indicator of ornament quality. Of 78 birds collected between 1992 and 1997, 42% carried helminths, including four species of digenean trematodes (flukes), three species of cestodes (tapeworms) and an acanthocephalan (spiny-headed worm). Faecal samples examined for helminth eggs in another 92 birds in 1998 and 2000 showed similar rates of infestation. Actively moulting bar-tailed godwits were confirmed to be heavier and to show more extensive breeding plumage than non-moulting birds. In females, but not in males, active moult was associated with fewer cestodes and acanthocephalans. Also, breeding plumage and presence of cestodes were negatively associated in females. We argue that the quality of the breeding plumage reliably indicates parasite resistance in female godwits. The repeatability of plumage scores of females between years is consistent with such resistance having a heritable component. In contrast, male ornaments may demonstrate other qualities, e.g. an ability to combine adequate fuelling and flight performances with moult during the time-stress of migration.     

Seasonal colour change by moult or by the abrasion of feather tips: a comparative study

Many birds undergo seasonal changes in plumage coloration by prebreeding moult, abrasion of cryptic feather tips, or both. Seasonal dichromatism is thought to result from optimizing coloration to the conflicting demands of different life-cycle periods, sexual selection for conspicuousness being substantial during the mating season, whereas selection for camouflage and for social signals may act in all seasons. Furthermore, energetic and time demands may constrain the extent of moult, thereby limiting colour change. We investigated the relative importance of several factors in shaping this variation in a songbird clade using phylogenetic comparative methods. We found that prebreeding moult relates most strongly to breeding onset and winter diet, demonstrating that both time and food availability constrain feather replacement. Feather abrasion was best predicted by winter flocking behaviour, and secondarily by open habitats, implying that exposure to predators and the simultaneous need for social signalling may favour the expression of partially obscured ornaments in the non-breeding season. The combined occurrence of prebreeding moult and feather abrasion was associated with the polygynous mating system, suggesting that species under strong sexual selection may employ both strategies of colour change to ensure the full expression of breeding coloration.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 711–721.     

Plasticity in moult speed and timing in an arctic‐nesting goose species

Environmental constraints are strong in migratory species that breed in the Arctic. In addition to breeding, Anatidae have to renew all their flight feathers during the short arctic summer. We examine how temporal constraints and climate affect the phenology of flight feather moult in the greater snow goose Chen caerulescens atlantica, a High Arctic nesting species. We used a database of 1412 moulting adult females measured over 15 yr on Bylot Island, Nunavut. Ninth (9th) primary length was used to determine the moult stage and speed of feather growth. We found a positive relationship between median annual hatching and moult initiation dates and the slope did not differ from 1. The interval between hatching and moult initiation was thus rather fixed and geese did not initiate moult earlier when reproductive phenology was delayed. Nonetheless, there was no relationship between median hatching date and the date at which birds regained flight capacity, suggesting that date of end of moult is independent of the reproductive phenology. There was a trend for an increase in the speed of flight feather growth in years with delayed hatching date. This is the most likely mechanism that could explain moult phenology adjustment in this species. Finally, we found a positive relationship between 9th primary length (corrected for inter‐annual variations) and body condition, suggesting a delay in moulting for individuals in poor condition. These results suggest that moult plasticity is primarily governed by variations in feather growth speed. This phenotypic plasticity could be necessary to complete flight feather renewal before the end of the arctic summer, independently of reproductive phenology and spring environmental conditions. Our novel results suggest possible phenological adjustments through moult speed, which was considered constant in geese until now.     

Continent-wide variation in feather colour of a migratory songbird in relation to body condition and moulting locality

Understanding the causes of variation in feather colour in free-living migratory birds has been challenging owing to our inability to track individuals during the moulting period when colours are acquired. Using stable-hydrogen isotopes to estimate moulting locality, we show that the carotenoid-based yellow-orange colour of American redstart (Setophaga ruticilla) tail feathers sampled on the wintering grounds in Central America and the Caribbean is related to the location where feathers were grown the previous season across North America. Males that moulted at southerly latitudes were more likely to grow yellowish feathers compared with males that moulted more orange-red feathers further north. Independent samples obtained on both the breeding and the wintering grounds showed that red chroma-an index of carotenoid content-was not related to the mean daily feather growth rate, suggesting that condition during moult did not influence feather colour. Thus, our results support the hypothesis that feather colour is influenced by ecological conditions at the locations where the birds moulted. We suggest that these colour signals may be influenced by geographical variation in diet related to the availability of carotenoids.     

A sexual conflict in collared flycatchers,Ficedula albicollis: early male moult reduces female fitness

A sexual conflict over levels of parental care occurs in most animals with biparental care, and studies of sexual differences in levels of parental care have usually focused on its intra-annual fitness consequences. We investigated inter-annual fitness consequences of a sexual difference in timing of feather replacement (moult) in collared flycatchers (Ficedula albicollis). In this study, males overlapped reproduction and moult more often than females, they also initiated their moult at an earlier stage of breeding than females. Females mated to males with a moult-breeding overlap had significantly lowered survival chances than females mated with males initiating moult after breeding. Furthermore, females mated with moulting males risked a lowered future fecundity in terms of a delayed start to breeding in the following season. However, early moulting males achieved a similar reproductive success as males initiating moult after breeding. Likewise, male survival probability to the following breeding season did not differ between early and late moulting individuals, nor was there any evidence that males gained or lost in future mating advantages by moulting early. These results show not only that a sexual conflict over timing of moult may operate, but also that it can impose severe fitness consequences, in terms of reduced future fecundity and survival probability, upon the ''losing'' sex.     

Spring migration of the Common Gull in Britain and Ireland

Due to the Collared Dove's extended breeding season, moulting birds may be found in all months. Young birds from late broods often arrest their post-juvenile moult over the midwinter period, and these birds may moult again later in the same year, in line with the moult schedule for adults.     

Conditions at the breeding grounds and migration strategy shape different moult patterns of two populations of Eurasian golden plover Pluvialis apricaria

Events in the life cycle of migrant birds are generally time‐constrained. Moult, together with breeding and migration, is the most energetically demanding annual cycle stages, but it is the only stage that can be scheduled at different times of the year. However, it is still not fully understood what factors determine this scheduling. We compare the timing of primary feather moult in relation to breeding and migration between two populations of Eurasian golden plover Pluvialis apricaria, the continental population breeding in Scandinavia and in N Russia that migrates to the Netherlands and southern Europe, and the Icelandic population that migrates mainly to Ireland and western UK. Moult was studied at the breeding grounds (N Sweden, N Russia, Iceland) and at stopover and wintering sites (S Sweden, the Netherlands). In both populations, primary moult overlapped with incubation and chick rearing, and females started on average 9 d later than males. Icelandic plovers overlapped moult with incubation to a larger extent and stayed in the breeding grounds until primary moult was completed. In contrast, continental birds only moulted the first 5–7 primaries at the breeding grounds and completed moult in stopover and wintering areas, such as S Sweden and the Netherlands. This overlap, although rare in birds, can be understood from an annual cycle perspective. Icelandic plovers presumably need to initiate moult early in the season to be able to complete it at the breeding grounds. The latter is not possible for continental plovers as their breeding season is much shorter due to a harsher climate. Additionally, for this population, moulting all the primaries at the stopover/wintering site is also not possible as too little time would remain to prepare for cold‐spell movements. We conclude that environmental conditions and migration strategy affect the annual scheduling of primary feather moult in the Eurasian golden plover.     

Moult of three Palaearctic migrants in their West African winter quarters

Some theories about moult strategies of Palaearctic passerine migrants assume that birds adapt timing of moult to environmental conditions such as rainfall on their African wintering grounds. Species wintering in the northern tropics should limit moult to the period shortly after their arrival at the end of the rainy season. Passerine migrants wintering in West Africa should also moult more rapidly compared to related species or conspecific populations that moult elsewhere. We investigated the moult of melodious warblers Hippolais polyglotta, willow warblers Phylloscopus trochilus and pied flycatchers Ficedula hypoleuca wintering in Comoé National Park, Ivory Coast, between October 1994 and April 1998. In contrast to previous studies we did not restrict our analyses to moult of flight feathers but also included moult of body feathers. The results differed partially from the general assumptions of previous authors. Melodious warblers moulted twice: a complete moult shortly after their arrival, and a moult of body feathers and in some cases some tertials and secondaries in spring. Willow warblers moulting flight feathers were found between December and March with the majority moulting in January and February. Primary moult was not faster compared to populations moulting in central Africa and South Africa. Body feather moult varied strongly among individuals with birds in heavy moult between December and April. Pied flycatchers moulted body feathers and tertials between January and April. Birds with growing feathers were found throughout the whole period including the entire dry season. Moult strategies are thus not readily related to a few environmental factors in general and our results show that factors other than mere resource availability during certain times on the wintering grounds are likely to govern the timing of moult.Communicated by F.Bairlein     

The effects of delaying the start of moult on the duration of moult, primary feather growth rates and feather mass in Common Starlings Sturnus vulgaris

In many species of birds there is a close relationship between the end of breeding and the start of moult. Late-breeding birds therefore often start to moult late, but then moult more rapidly. This is an adaptive mechanism mediated by decreasing day lengths that allows late-breeding birds to complete moult in time. This study asked how these birds complete moult of the primary feathers more rapidly, and the consequences of this on the mass of primary feathers. Common Starlings Sturnus vulgaris were induced to moult rapidly in one of two ways. In the first experiment, one group was exposed to artificially decreasing photoperiods from the start of moult, whereas the control group remained on a constant long photoperiod. The second experiment was a more realistic simulation. Two groups were allowed to moult in an outdoor aviary. One group started to moult at the normal time. In the other, the start of moult was delayed by 3 weeks with an implant of testosterone. The duration of moult was significantly reduced in both the group experiencing artificially decreasing photoperiods and the group in which the start of moult was delayed. The faster moult rate was achieved by moulting more feathers concurrently. The rate of increase in length of each of the primary feathers, and their final length, did not differ between groups. The rate at which total new primary feather mass was accumulated was greater in more rapidly moulting birds, but this was insufficient to compensate for the greater numbers of feathers being grown concurrently. Consequently, the rate of increase in mass of individual feathers, and the final feather mass, were less in the rapidly moulting birds. A 3-week delay in the start of moult is not an unrealistic scenario. That this caused a measurable decrease in feather mass suggests that late-breeding birds are indeed likely to suffer a real decrease in the quality of plumage grown during the subsequent moult.     

THE PRIMARY MOULT OF WADERS ON THE ATLANTIC COAST OF MOROCCO

Data from 3659 waders of 23 species live-trapped in the years 1971-73 on the Atlantic coast of Morocco during the period of autumn moult and migration are analysed to estimate duration and timing of primary moult. Common Sandpiper was the only species to moult primaries in its first autumn (unless published ageing criteria are incorrect). Several species showed a low incidence of arrested primary moult and a higher incidence was observed in Ringed, Kentish and Grey Plovers. This is discussed in relation to breeding and migration. Similar rates of primary feather replacement relative to specific moult duration were observed in all species for which information was available. Comparisons between species and with published studies showed that variations in rate of moulting between species and between different geographical populations of the same species were largely due to differences in feather growth rate rather than in the numbers of primaries concurrently in growth. Variations in rate between individuals of the same population were achieved, at least in the first part of moult, by differences in feather dropping rate resulting in differences in the numbers of primaries growing concurrently. The timing and duration of moult in different populations and differences between breeding and non-breeding components were closely related to the requirements of other annual cycle activities, notably breeding and migration. Non-breeding birds summering in Morocco had started moult early. Locally breeding birds had an early start to a fairly slow moult which overlapped with breeding and which in some cases passed through an arrested stage. Birds breeding in cold temperate and arctic regions and wintering in Morocco moulted in a short time soon after arrival. In some cases, notably in Ringed Plovers, birds had commenced moulting on the breeding grounds and arrested moult during migration. Most Redshank and possibly Dunlin migrated in active wing moult. The fastest primary moult was achieved by high arctic breeding birds, Curlew Sandpiper and possibly Little Stint, which stopped to moult in Morocco before moving on to wintering areas further south. This situation is contrasted with that of populations of these two and other species wintering in the southern hemisphere where moult occurs over an extended period during the northern winter.     

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla . As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 98–105.     

Resistance of flight feathers to mechanical fatigue covaries with moult strategy in two warbler species

Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.     

THE MOULT OF THE BULLFINCH PYRRHULA PYRRHULA

The distribution of feather tracts and their sequence of moult in the Bullfinch is described. The adult post-nuptial moult, which is complete, lasted 10–12 weeks, and the post-juvenile moult, which is partial, 7–9 weeks. Adult moult began with the shedding of the first (innermost) primary and ended with the replacement of the last. Variations in the rate of moult in the flight feathers were mainly achieved, not by changes in the growth rates of individual feathers, but in the number of feathers growing concurrently. The primaries were shed more slowly, and the onset of body moult delayed, in birds which were still feeding late young. In 1962, the onset of moult in the adults was spread over 11 weeks from thc end of July to the beginning of October, and in the two following years over the six weeks, from the end of July to the beginning of September. The onset of moult was delayed by late breeding, which itself occurred in response to a comparative abundance of food in late summer, markedly in 1962. In all years, the first juveniles to moult started at the end of July, and the last, three weeks after the latest adults. Juveniles moulting late in the season retained more juvenile feathers than those moulting earlier. During moult, adult and juvenile Bullfinches produce feathers equivalent to 40% and 33% respectively of their dry weights. In both, for much of the moult, an average of nearly 40 mgm. of feather material—some 0.6% of their dry-weight–is laid down each day. The remiges of the adult comprise only a seventh of the weight of the entire plumage, and it is suggested that their protracted moult results not so much from their energy requirements, as from the need to maintain efficient flight. Variation in the rate of moult in the remiges was much less pronounced than in the body feathers. Bullfinches were less active during moult than at other times of the year. The weights of both adults and juveniles increased during moult. The food during the moult period is described. In all years, most Bullfinches finished moulting just before food became scarce, even though this occurred at different times in different years. In one year, adults moulting latest in the season probably survived less well than those moulting earlier; the same was apparently true of the juveniles in all years. The timing of moult in the Bullfinch, and the factors initiating it, are discussed in relation to the breeding season and foodsupply near Oxford.     

两种山鹪莺的春季换羽及尾羽变化

换羽是鸟类为保证持续生存的重要过程。换羽策略与鸟类进化及对环境的适应紧密相关,研究鸟类换羽特征,对于了解鸟类的分类、系统发育、进化历史及其对环境的适应性等方面都有重要意义。2007年3月至9月,在广东肇庆市江溪村对黄腹山鹪莺(Prinia flaviventris)和纯色山鹪莺(P.inornata)的春季换羽进行了研究。通过设置雾网捕捉2种山鹪莺,对捕捉到的成体进行体重及身体量度的测量;对飞羽及尾羽进行标记:初级飞羽以翅尖的第一枚羽毛标记为"P1",次级飞羽以翅中部最外一枚标记为"S1",向内依次递增标记;尾羽以中央两根最长尾羽为"T1",分别向两侧递增标记为"T2~T5"。采用单因素方差分析(One way ANOVA)对不同月份山鹪莺的体重值进行差异性检验,对体重与月份进行Pearson相关分析,对尾羽的长度和宽度进行Pearson偏相关分析(控制变量:体长)。研究结果表明:1)两种山鹪莺换羽期为3至5月,持续时间约为60 d;2)两种山鹪莺春季换羽仅更换尾羽,换羽模式均为离心型,即中央一对尾羽最先开始替换,然后向两侧由内到外逐次更替;3)两种山鹪莺的尾羽长度和宽度同步变化,但绝大部分山鹪莺非繁殖期尾羽长度与繁殖期尾羽长度之比大于非繁殖期尾羽宽度与繁殖期尾羽宽度之比,即繁殖期尾羽相对较宽;4)两种山鹪莺换羽期间体重大致呈现下降趋势,但变化不显著(P0.05)。推测两种山鹪莺通过增加食物的摄入来抵抗换羽期和繁殖期重叠而导致的能量消耗,这可能与该地区丰富的食物资源有关,并在一定程度上体现了两种山鹪莺换羽策略对环境的适应性。     

Yellow plumage colour of Great Tits Parus major correlates with changing temperature and precipitation

Weather is known to affect the phenology and behaviour of birds, but weather-related changes of phenotypic traits involved in communication have received little attention. Using an 8-year dataset, we investigated links between carotenoid-based reflectance of the freshly moulted breast of Great Tits Parus major, weather during the moulting period and food availability during the preceding breeding season, and we investigated interannual changes. In both sexes, we found a change of colour expression to more saturated and darker yellow over the study period in parallel with increasingly dry and warm weather during moult. These results indicate that the expression of traits playing roles in communication may be controlled by weather and may therefore shift in response to changing climate.     

Wing moult in relation to autumn migration in adult Common Whitethroats Sylvia communis communis

Most long-distance passerine migrants in Sweden moult on breeding grounds before leaving on autumn migration to winter quarters. However, birds laying second or replacement clutches, or just breeding late, have too little time for a normal moult on the breeding grounds. When time is limited the birds may respond by making various adjustments to the moult, for example by moulting more quickly or by suspending the moult. In this study, the relationship between the performance of post-nuptial remex moult in Common Whitethroats breeding on Gotland, southeast Sweden, and autumn migration departure was investigated. The majority (77%) of the birds had interrupted moult in either the primaries or secondaries. Interruption of moult was more common among birds with a later onset date, as was asymmetry in moult between wings. The interruption of moult led to a significant time gain and moult completion was, consequently, more synchronized than moult onset. The results from this study indicate, in accordance with other data, that an early start of autumn migration is important. An early start may be crucial to facilitate the crossing of the Sahara Desert once the dry season has begun.     

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.     

Sexual differences in moult–breeding overlap and female reproductive costs in pied flycatchers, Ficedula hypoleuca

1. In this study I show that a sexual difference in timing of the post-nuptial moult frequently occurs in a sub-arctic population of the pied flycatcher.
2. Most pairs started to moult after fledging of their young, but an overlap between moult and nestling feeding was more common among males than females. This sexual difference in moult–breeding overlap increased as the season progressed.
3. Females with moult–breeding overlap laid smaller clutches than non-moulting females. In addition to many other factors explaining the seasonal decline in clutch size that has been found for many bird species, it is possible that females adjust their clutch size according to their own risk of having to start moulting while still feeding the nestlings.
4. Nearly 24% of the females were deserted by their mate before the young fledged. Desertion imposed no fitness costs to males in terms of fledgling number or quality, suggesting that their females managed to adjust their care for the loss of male care.
5. Deserted females started moulting later than aided females, which may be a result of their increased reproductive investment.
6. Deserted females and females aided by moulting males had lower survival rate than females aided by non-moulting males.
7. These findings suggests that delayed moult may be one mechanism causing inter-annual reproductive costs in birds, and the relationship between a sexual difference in timing of moult and its fitness consequences may be widespread among passerine birds.