The effects of sex, age and breeding success on breeding dispersal of pied flycatchers along a pollution gradient

We modelled breeding dispersal of an insectivorous bird, the pied flycatcher (Ficedula hypoleuca) around a point source of heavy metals (a copper smelter). We tested for the effects of sex, age, breeding success and environmental pollution on breeding dispersal distances of F. hypoleuca females and males. Unlike many earlier studies on breeding dispersal, we took into account distance-dependent sampling bias by including in our model the recapture probabilities at different distances from the site of origin. Our results show that F. hypoleuca females disperse much farther (on average 7.9 km) from their breeding sites than what was previously thought. In contrast, males only disperse short distances (on average 190 m). Breeding success affected female breeding dispersal distances depending on female age: young females moved on average 8 km from their previous breeding place irrespective of their breeding success, while old females only seemed to move this far when their fledgling production was good. Females successful in their breeding dispersed as far as less successful females, or, among old birds, even farther. Females which dispersed long distances produced more fledglings after the movement than those staying near their previous breeding site. Degree of environmental pollution had no effect on female or male breeding dispersal distances. A polluted and unproductive environment does not seem to stimulate F. hypoleuca parents to move to more profitable territories.     

Pollution related effects on immune function and stress in a free-living population of pied flycatcher Ficedula hypoleuca

We investigated whether exposure to heavy metal pollution affected the immune function of individuals in a free living population of a small insectivorous passerine bird, the pied flycatcher Ficedula hypoleuca . We measured humoral immune responses in two study areas: a polluted area in the vicinity of a copper smelter and a control area far from the smelter. Plasma corticosterone level and blood heterophil/lymphocyte ratio (H/L) were used as more general physiological measures of stress. The immune response of F. hypoleuca was not suppressed by pollution stress. In contrast, we found that F. hypoleuca males showed stronger humoral immune responses to a novel antigen (tetanus toxoid) in the polluted environment than in the unpolluted one. After the immunization of males, numbers of lymphocytes rose significantly more in the polluted area, leading to a smaller H/L ratio than in males from the control area. Females showed no pollution related effects on their immune responses. Corticosterone levels of males and nestlings were not related to pollution levels. Nestlings showed somewhat higher H/L ratios and lower fledging success in the polluted area, both factors indicating increased stress levels in a polluted area. Our results suggest that humoral immune response of male F. hypoleuca may be enhanced under moderate levels of heavy metal pollution. Enhanced immune function may, however, also be costly for birds and the higher humoral immune responses in polluted areas may thus have negative effects on the birds' breeding performance and survival.     

Environmental pollution affects genetic diversity in wild bird populations

Many common environmental pollutants, together with nuclear radiation, are recognized as genotoxic. There is, however, very little information on pollution-related genetic effects on free-living animal populations, especially in terrestrial ecosystems. We investigated whether genetic diversity in two small insectivorous passerines, the great tit (Parus major) and the pied flycatcher (Ficedula hypoleuca), was changed near point sources of heavy metals (two copper smelters) or radioactive isotopes (nuclear material reprocessing plant). We measured concentration of heavy metals and nucleotide diversity in mitochondrial DNA in feather samples taken from nestlings in multiple polluted areas and at control sites. In both species, heavy metal concentrations - especially of arsenic - were increased in feathers collected at smelter sites. The P. major population living near a smelter showed significantly higher nucleotide diversity than a control population in an unpolluted site, suggesting increased mutation rates in a polluted environment. On the contrary, F. hypoleuca showed reduced nucleotide diversity at both smelter sites but increased nucleotide diversity near the source of radioactivity. Our results show that heavy metal pollution and low level nuclear radiation affect the nucleotide diversity in two free-living insectivorous passerines. We suggest that the different response in these two species may be due to their different ability to handle toxic compounds in the body.     

Social environment affects juvenile dispersal in great tits (Parus major)

1.?Habitat selection can affect individual fitness, and therefore, individuals are expected to assess habitat quality of potential breeding sites before settlement. 2.?We investigated the role of social environment on juvenile dispersal behaviour in the great tit (Parus major). Two main contradictory hypotheses can be formulated regarding social effects on juvenile dispersal as follows: (i) High fledgling density and sex ratio may enhance the intensity of local (kin) competition and, therefore, reduce individual survival chance, enhance emigration and reduce settlement ('repulsion' hypothesis) (ii) Alternatively, high fledgling density and sex ratio may signal high-quality habitat or lead to aggregation and thus increase individual survival chance, reduce emigration and enhance settlement ('attraction' hypothesis). 3.?To disentangle positive from negative effects of high density and male-biased sex ratio on dispersal, we manipulated the social composition of the fledgling population in 12 semi-isolated nest-box areas (plots) via a change of fledgling density (low/high) as well as fledgling sex ratio (female-biased/balanced/male-biased) across 3?years. We then tested whether experimental variation in male and female fledgling densities affected variation in local survival, emigration and settlement of juveniles, and whether social effects on survival and dispersal support the 'repulsion' or 'attraction' hypothesis. 4.?We found no experimental effects on local survival and emigration probabilities. However, consistent with the 'attraction' hypothesis, settlement was significantly and positively affected by local experimental sex ratio in each of the study years: both male and female juveniles avoided female-biased plots and settled more in plots that were balanced and male-biased the previous year. 5.?Our study provides unprecedented experimental evidence that local sex ratio plays a causal role in habitat selection. We suggest that settlers avoid female-biased plots because a high proportion of females may reflect the absence or the low quality of local resources in the habitat. Alternatively, male territory acquisition may be facilitated by a high local density of 'candidate' males, and therefore, juveniles were less successful in settling in female-biased plots.     

Variation in the hematocrit of a passerine bird across life stages is mainly of environmental origin

The heritability of the hematocrit in adult, breeding pied flycatchers Ficedula hypoleuca is examined in a southern European population across seven years to see the consistency, or lack thereof, of patterns found with the trait at fledgling age where no significant heritability could be detected. While the across-years repeatability of the trait in adult, breeding birds was low but significant, heritabilities based on adult parent-adult offspring regressions controlling for assortative mating and full-sibling comparisons did not differ significantly from zero. Neither were heritability estimates affected by selection on fledgling hematocrit, as it was unrelated to local recruitment. There was no relationship between fledgling and adult hematocrit, suggesting that both behave as phenotypically plastic, different traits. This is the first study testing for the heritability of a physiological trait in birds with data from adult, free-ranging individuals.     

Lifetime fitness of short- and long-distance dispersing great reed warblers

Dispersal is of prime importance for many evolutionary processes and has been studied for decades. The reproductive consequences of dispersal have proven difficult to study, simply because it is difficult to keep track of dispersing individuals. In most previous studies evaluating the fitness effects of dispersal, immigrants at a study locality have been lumped into one category and compared to philopatric individuals. This is unfortunate, because there are reasons to believe that immigrants with long and short dispersal distances may differ substantially in reproductive success. In the present study, we used a combination of capture-recapturing and multilocus microsatellite genotyping to categorize great reed warblers at our Swedish study site as philopatric individuals or short- or long-distance dispersing immigrants. We then performed novel comparisons of lifetime reproductive success (LRS) and survival rates of these three dispersal categories. The birds belonged to cohorts 1987-1996, and data for their LRS were gathered between 1988 and 2003. The analyses showed that philopatric males attracted more females, produced more fledglings and recruits throughout their lives, and survived better than immigrants. Among the immigrant males, those categorized as long-distance dispersers had lowest LRS and survival probability. Models that included covariates of potential importance showed that the difference in LRS between dispersal categories was partly caused by corresponding variation in number of breeding years at our study site. These results indicate that short- and, in particular, long-distance dispersers were of poor phenotypic quality, but it may also be proposed that immigrants attracted few females because they were poorly adapted to the local social environment. In females, the number of local recruits corrected for the number of breeding years (as well as for number of fledglings) differed between dispersal categories in a pattern that suggests an intermediate optimal dispersal distance. Short-distance dispersers recruited more offspring per year (and per fledgling) than both philopatric individuals and long-distance dispersers. Data suggest that the low LRS of philopatric females was related to costs of inbreeding. The low LRS of long-distance dispersing females may have resulted from their offspring being especially prone to disperse outside the study area, but also other potential explanations exist, such as local maladaptation. Our study highlights the importance of separating immigrant birds on the basis of their genetic similarity to the local study population when analyzing variation in LRS and inferring realized gene flow.     

Manipulation of offspring sex ratio by second-mated female house wrens

In 1973, Trivers and Willard proposed that offspring sex ratio should be associated with the quality of parental care likely to be provided to the offspring. We tested this hypothesis by comparing fledgling sex ratios in nests of first- and second-mated female house wrens (Troglodytes aedon). In our Wyoming population, second-mated females typically receive little or no male parental assistance and fledge fewer and lower-quality young compared with first-mated females. Assuming that being of lower quality has stronger negative effects on the future reproductive success of males than that of females in this polygynous population, we predicted that fledgling sex ratios in the nests of second-mated females would be female-biased compared with the fledgling sex ratios of first-mated females. Additionally, we asked whether any sex bias at fledging could have resulted from male-biased nestling mortality caused by sex-biased parental provisioning. As predicted, mean fledgling sex ratios in nests of second-mated females were more female-biased than fledgling sex ratios in nests of first-mated females. However, we found no evidence of either sex-biased nestling mortality or sex-biased parental provisioning. These findings suggest that females are responding to their status as second-mated females and to the associated low-quality parental care that their young are likely to receive by producing female-biased clutches rather than manipulating the offspring sex ratio through sex-biased nestling mortality.     

Reed bunting females increase fitness through extra-pair mating with genetically dissimilar males

Females of many socially monogamous species accept or even actively seek copulations outside the social pair bond. As females cannot increase the number of offspring with promiscuous behaviour, the question arises why they engage in extra-pair mating. We used microsatellite data to determine paternity, heterozygosity and genetic relatedness in the reed bunting (Emberiza schoeniclus), a species with high levels of extra-pair paternity (EPP). We found that extra-pair young (EPY) were more heterozygous than within-pair young (WPY). The high heterozygosity of the EPY resulted from a low genetic similarity between females and their extra-pair mates. EPY were heavier and larger when compared with their maternal half-siblings shortly before they left the nest. Recapture data indicated a higher fledgling survival of EPY compared with WPY. Our data suggest that reed bunting females increase the viability of their offspring and thus fitness through extra-pair mating with genetically dissimilar males.     

Polygyny and its fitness consequences for primary and secondary female pied flycatchers

In polygynous species with biparental care, the amount of paternal support often varies considerably. In the pied flycatcher (Ficedula hypoleuca), females mated with monogamous males receive more male assistance during the nestling phase than females mated with bigynous males, as the latter have to share their mates with another female. Bigynous males, however, give more support to their primary broods than to their secondary broods. Using a long-term dataset (31 years), the present study revealed that direct reproductive success, i.e. number of fledglings, was lower in females that mated with bigynous males, especially in secondary broods without male assistance, than in females that mated with monogamous males. Secondary broods with male assistance were more affected than primary broods. Female survival was independent of mating status. In primary broods, a delayed compensation for inferior direct reproductive success was found in terms of the number of grandoffspring, a phenomenon that did not occur in secondary broods. Delayed compensation in primary broods refers to indirect effects, i.e. good genes. According to the sexy son hypothesis, genetically superior (i.e. sexy) males may have sons with a higher number of broods belonging to a polygynous breeding status than do sons from broods with a monogamous father. This was indeed the case for sons descending from primary broods, but not for sons descending from secondary broods.     

Behavioural and Genetic Data Suggest that Bechstein's Bats Predominantly Mate Outside the Breeding Habitat

Investigating mating systems of species with a cryptic lifestyle often requires a combination of behavioural and genetic data. We used such a combined approach to investigate the mating system of the communal breeding Bechstein's bat (Myotis bechsteinii). Although females of this species are philopatric, gene flow among colonies is high. Gene flow occurs if dispersing males mate within the colony to which they moved. Males could gain local matings by defending resources or females in their breeding habitat. Alternatively, mating may take place at swarming sites, apart from the breeding habitat of the females, where males and females of several colonies meet. Whether or not males defend resources or females in the breeding habitat is of importance for understanding the mating system. Detailed observations of individual foraging and roosting behaviour over 4 yr suggest that males do not defend resources or females to gain matings. Moreover, paternity assignment based on microsatellite data of four complete juvenile cohorts showed that local males fathered less than 25% of the juveniles born in the colony where they settled. Even more striking, none of the males that had immigrated into our study area reproduced with the local females.     

Gender and viability selection on morphology in fledgling pied flycatchers

Until recently, analyses of gender-dependent differences in viability selection and the ontogeny of sexual size dimorphism have been plagued by difficulties in determining the sex of nestling birds on the basis of morphology. Recently, this problem was overcome using molecular sex identification to report for the first time body-size-mediated antagonistic selection on the viability of male and female collared flycatchers. We used molecular sex identification to analyse natural selection on fledgling viability, sexual size dimorphism and effects of parasites in relation to gender in a Mediterranean population of the related pied flycatcher Ficedula hypoleuca. There was directional positive selection on fledgling weight but no selection on tarsus length. Fledgling weight was the most important determinant of fledgling survival, with heavier fledglings having increased viability. Although selective trends were of the same sign for both sexes, only among female fledglings were selection differentials and gradients statistically significant. Therefore, similar trends in selection were revealed in analyses of a data set where sex was ignored and in separate analyses using same-sex sibship trait means. Mite nest ectoparasites negatively affected fledgling weight, and the effects were stronger in female than male fledglings. There was no effect of parasitism on the tarsus length in males, as previously reported in retrospective analyses performed without knowledge of sex until recruitment. Overall, selection on fledgling viability on the basis of morphological traits and hatching date was not confounded by an individual's gender.     

Population and social dynamics changes in ring-tailed lemur troops at Berenty,Madagascar between 1989 - 1999

In the present study, we recorded all births, immigrations, deaths, and emigrations for a population of ring-tailed lemurs at Berenty Reserve, Madagascar, between September 1989 and August 1999. In September 1989, three troops (C, B, and T) inhabited the study area of 14.2 ha. During the 10-year period, eight troop divisions, six evictions of females, and three troop takeovers of ranges by other troops occurred in and around the study area. Consequently, in August 1999, the number of troops in the same area increased to six (CX, C1, C2A, C2B, T1, and T2). The number of lemurs aged >1 year increased from 63 to 82, which resulted from 204 births, 58 immigrations, 125 deaths, and 118 emigrations. Of the 204 newborn lemurs during the study period, 103 died, 44 emigrated outside the study area, and 57 remained within the study area. The total number of lemurs that emigrated from natal troops was 69 (54 males and 15 females). Natal males left their troops around the age of 3. Non-natal males changed troops after a tenure varying from 1 to 7 years. Survival curves showed a fall in survival rates of both sexes to < 0.5 between the ages of 2 and 3. For females, the survival rate gradually decreased to < 0.2 at the age of 9. On the other hand, due to emigration, the survival rate of males could not be determined after the age of 5 yr. Since some males attained high-rank at the age of 6 – 10 yr, the prime age for male ring-tailed lemurs is thought to be around 7 – 10 yr. Ring-tailed lemurs are essentially female philopatric, because all cases of females leaving natal troops resulted from troop divisions or forced evictions. Such social changes may have resulted from competition among females. All cases of troop divisions or evictions occurred in larger troops consisting of ≥20 lemurs, and only a few females could rejoin their troops. When males joined such a female-group, a new troop was formed. Although promoted by an increase in population, frequent emigrations of females from original troops are the characteristics of ring-tailed lemurs at Berenty.     

High survival rate and site fidelity in the Siberian Tit Parus cinctus, a focal species of the taiga

The Siberian Tit Parus cinctus population of Finland, and probably of the whole of Fennoscandia, has declined dramatically during this century. Understanding its population dynamics is essential for its conservation. We studied annual survival rate and dispersal distance of both breeding and fledgling Siberian Tits during 1989-97 in a moderately managed forest habitat near the southern border of its range in Kuusamo, northeastern Finland. Breeding density was low, averaging 0.51 pairs/km in a nestbox area of about 42 km². This was probably an underestimate, because we did not search for nests in natural holes. However, following its population decline, overall densities at the southern border of the range are low. The study area was, however, suitable breeding habitat as reflected by their high nesting success. Brood size at fledging averaged 7.45 young, and reproductive output was 6.16 young per breeding pair, including failed nests. Clutch size adjustment was successful among pairs producing fledglings because, on average, 92.6% of the eggs laid produced young in successful broods. Median distance between consecutive breeding attempts was 100 m for both males and females (range 0–2500 m and 0–6000 m, respectively). Natal dispersal distance was significantly longer in females than in males. We applied Cormack-Jolly-Seber modelling to estimate survival and recapture probabilities separately. Survival of breeding birds was not sex-related, averaging 0.69 annually. Some of the ringed fledglings (3.0%) were later captured as breeders in the area. This is an underestimate of local survival rate due to incomplete recapturing of breeding birds. The implications of these results with respect to the conservation status of the species are discussed.     

Patterns of breeding behaviour and plasma levels of hormones in a free-living population of Pied flycatchers, Ficedula hypoleuca

Plasma levels of luteinizing hormone (LH) and steroid hormones were measured by radioimmunoassay in a series of plasma samples collected from a free-living population of Pied flycatchers, Ficedula hypoleuca. Both males and females have similar circulating levels of LH and testosterone when they arrive in the breeding area. However, as territories and pair bonds are established, and as courtship and nest-building ensue, the plasma levels of testosterone in males become elevated over those of females, although a lesser peak also occurs in females. As the egg laying phase and incubation begin, plasma levels of LH and testosterone decline sharply in both sexes. On the other hand polyterritorial males, that establish secondary territories, maintain higher levels of LH and testosterone than males remaining in the home-territory, at least until they return to the home-territory, usually at the middle part of the incubation period. By this time polyterritorial males also have basal levels of these hormones. Curiously, females on home-territories show a small, but significant, rise of circulating LH and testosterone at the time the males return.
In males, plasma levels of corticosterone are elevated from the time of arrival on the breeding grounds until the end of the nest-building period. Thereafter corticosterone levels decline in males staying on the home-territory but remain elevated in polyterritorial males until the latter return to the home-territory. In contrast, circulating levels of corticosterone in females are only elevated for brief period during egg laying.     

Mate choice in Pied Flycatchers Ficedula hypoleuca: can females use song to find high‐quality males and territories?

High-quality male Pied Flycatchers Ficedula hypoleuca (defined by brighter plumage, better condition and more experience) have previously been shown to have larger syllable repertoires and greater song versatility than males of inferior quality. Thus, by preferring more complex songs, females could choose a high-quality male. Females may also use song as a cue to find a high-quality territory since early arriving males may obtain the best territories and these males have more complex songs than late-arriving males. We found that males with more complex songs had a greater chance of becoming paired and stayed unpaired for a shorter period than males with less elaborate songs. When controlling for arrival order, however, only strophe versatility was still correlated with pairing order. Males defending popular territories had more complex and longer songs and were also in better body condition than males in less popular territories. A multiple logistic regression showed that song length was important in explaining whether a male defended a popular nestbox or not. Thus, male arrival time seems to be important in deciding the quality of a male's territory, which in turn explains female choice. However, song quality seems to add important information. Thus, females could find both high-quality males and high-quality territories by using song cues during mate choice.     

Survival during the Breeding Season: Nest Stage,Parental Sex,and Season Advancement Affect Reed Warbler Survival

Avian annual survival has received much attention, yet little is known about seasonal patterns in survival, especially of migratory passerines. In order to evaluate survival rates and timing of mortality within the breeding season of adult reed warblers (Acrocephalus scirpaceus), mark-recapture data were collected in southwest Poland, between 2006 and 2012. A total of 612 individuals (304 females and 308 males) were monitored throughout the entire breeding season, and their capture-recapture histories were used to model survival rates. Males showed higher survival during the breeding season (0.985, 95% CI: 0.941–0.996) than females (0.869, 95% CI: 0.727–0.937). Survival rates of females declined with the progression of the breeding season (from May to August), while males showed constant survival during this period. We also found a clear pattern within the female (but not male) nesting cycle: survival was significantly lower during the laying, incubation, and nestling periods (0.934, 95% CI: 0.898–0.958), when birds spent much time on the nest, compared to the nest building and fledgling periods (1.000, 95% CI: 1.00–1.000), when we did not record any female mortality. These data (coupled with some direct evidence, like bird corpses or blood remains found next to/on the nest) may suggest that the main cause of adult mortality was on-nest predation. The calculated survival rates for both sexes during the breeding season were high compared to annual rates reported for this species, suggesting that a majority of mortality occurs at other times of the year, during migration or wintering. These results have implications for understanding survival variation within the reproductive period as well as general trends of avian mortality.     

Determinants of parental care and offspring survival during the post-fledging period: males care more in a species with partially reversed sex roles

Sexual conflict is magnified during the post-fledging period of birds when the sexes face different trade-offs between continuing parental care or investing in self maintenance or other mating opportunities. Species with reversed sex roles provide a unique opportunity to study the relationship between mating systems and investment in parental care. Here, we provide the first detailed study of the length of care by males versus females (n = 24 pairs) during the post-fledging period, assessing factors that may promote care within and between the sexes. In the northern flicker Colaptes auratus, a species with partly reversed sex roles, males cared longer than females (average 16 versus 12 days, respectively). Overall, 36 % of females but no males deserted the brood prior to fledgling independence. Parents that provisioned nestlings at a high rate also spent more days feeding fledglings. Among males, age and nestling feeding rates were positively associated with the length of care. Among females, a low level of feather corticosterone (CORT_f) was associated with a longer length of care. About 45 % of fledglings died within the first week, but fledglings with intermediate body mass had the highest survival suggesting stabilizing selection on mass. Fledgling survival was also higher in individuals with larger broods and lower levels of CORT_f. We demonstrate that because females can be polyandrous they often desert the brood before males, and that the sexes respond to different cues relating to their energy balance when deciding the length of care given to their offspring.     

Plasma prolactin concentrations in breeding pied flycatchers (Ficedula hypoleuca) with an experimentally prolonged brooding period

This experiment explored the stimulatory effect of brooding newly hatched young on plasma prolactin concentration in male and female pied flycatchers, Ficedula hypoleuca. By exchanging offspring between nestboxes, one group of parents was exposed to 1- to 3-day-old nestlings for 12 days. Females in this group continued night-time brooding during these 12 days. The females, but not the males (who do not participate in brooding nestlings), after 6 days had higher plasma prolactin concentrations than control birds which had shown a 50% reduction in night-time brooding at this stage. By Day 12, however, prolactin concentration had decreased to the same level as in control birds, even though these females were still brooding 3-day-old nestlings. Female flycatchers given 3-day-old nestlings on the day their own eggs hatched showed an earlier reduction in night-time brooding and an earlier decrease in plasma prolactin than did control birds. An early decrease in prolactin was also seen in the males of this experimental group.     

Behavioral, hormonal, and morphological responses of free-living male pied flycatchers to estradiol treatment of their mates

A field experiment was performed implanting female pied flycatchers (Ficedula hypoleuca) during the nest-building period with silastic tubes containing 17 beta-estradiol. Control females were given empty silastic tubes. The pied flycatcher is a polygamous and polyterritorial bird species. During two successive breeding seasons effects on reproductive behavior and the reproductive systems of males were studied. Reproductive success was not affected by the treatment. Estradiol treatment of females caused marked changes in the behavior of males, increasing aggressive and mate-guarding behavior. The results of the present study also support the idea that cuckoldry is a problem for the territory holder. Fewer males paired with estradiol-treated females established secondary territories, and significantly more of the polyterritorial males from the experimental population showed up in the home-territory when a simulated territorial intrusion was performed there. Control males were much more successful in getting a second female to breed. Spermatogenesis and hormonal cycles (DHT, LH, and testosterone) in males were not prolonged as a result of estradiol treatment of the females.     

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.