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.     

Annual variation in the timing of breeding and moulting in male and female Pied Flycatchers Ficedula hypoleuca

During five breeding seasons, the timing of breeding and moulting was studied in the Pied Flycatcher Ficedula hypoleuca. In central Sweden, on average 67% of the males and 41% of the females started moulting before the young fledged. The proportion of individuals with an overlap between breeding and moulting varied considerably between years, with the highest proportion of moulting males being recorded in the year when the females started egg-laying on the latest date. Despite a large annual variation in the proportion of individuals showing a moult/breeding overlap, the duration of this overlap varied insignificantly between years. The onset of moult in males seemed to be related to both calendar date and timing of the current breeding attempt. Most females postponed their moult until just before or just after the fledging of their young, independent of calendar date. There was no significant relationship between male and female moult scores and nestling weight at fledging or fledging success of their brood. Thus, in long-distance migrants such as Pied Flycatchers, it may be adaptive to have some overlap between reproduction and moult, but there seems to be a limit to how early in the breeding cycle they are able to start moulting.     

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.     

REVIEW

Mlingwa, C.O.F. 2000. Breeding and moult phenology of an avian community in the Pugu Hills, Tanzania. Ostrich 71 (1 &; 2): 87–90.

The breeding and moult phenology of an avian community in the Pugu Hills in coastal Tanzania was investigated in a ringing study carried out during January to March 1993, June 1994 to May 1995, and October 1995. A total of 74 bird species was examined. Breeding activity was concentrated from October to April with peak activity between November and February/March. Little or no breeding activity was recorded in May-September period. Moult, on the other hand, was much more spread throughout the annual cycle. Only at the end of the dry period in September and the start of the breeding season in October/November was moulting activity very low. These results are discussed in relation to seasonality of the environment in the study area. It is also suggested that although both processes are physiologically very costly, breeding is much more critical than moult hence must be timed to occur during the best period of the annual cycle.     

Moult strategies of Cory’s Shearwaters Calonectris diomedea borealis: the influence of colony location, sex and individual breeding status

The replacement of old feathers is essential for birds, but it is also an energy-demanding task. As moult usually does not coincide with other stressful events in its annual cycle, such as reproduction and migration, the bird can optimise its use of time and energy allocated to different activities. There are very few studies comparing the moult strategies of populations with similar breeding calendars but occurring in areas of different habitat quality. Cory’s Shearwaters Calonectris diomedea have a partial moult–breeding overlap, an unusual phenomenon among pelagic seabirds. We have compared the moult schedules in Cory’s Shearwater colonies located in distinct environments (pelagic vs. coastal) and show that moult–breeding overlap is less extensive on Selvagem Grande, situated in deep oceanic waters, than on Berlenga, situated on the continental shelf. Colony attendance of failed breeders, most of which were moulting, was lower at Selvagem Grande than at Berlenga, which suggests that the feeding areas of birds from the former site are more distant from the colony. Failed breeders started to moult earlier than individuals still raising a chick, and breeding status had a stronger influence on determining the onset of wing-feather moult than colony location. Despite published evidence that internal circannual rhythms and external cues, such as variation in daylength, are important factors regulating moult schedules, it is clear that birds retain a considerable flexibility that allows them to respond to external factors in order to strategically manage time and energy in a way that is thought to maximise their fitness.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

European Swallows Hirundo rustica in Botswana during three non-breeding seasons: the effects of rainfall on moult

van den Brink, B., Bijlsma, R.G. & van der Have, T.M. 2000. European swallows Hirundo rustica in Botswana during three non-breeding seasons: the effects of rainfall on moult. Ostrich: 71 (1): 198–204.

The rate of moult of European Swallows spending the non-breeding season in Botswana was studied during December-January of 1992/93,1993/94 and 1994/95 to investigate the effects of variability in rainfall and roosting habitat availability. In January 1994, 2–3 million European Swallows were counted at a traditional roost along the Boteti River. The rate of moult was relatively slow, about one feather (primary, secondary or tail feather) was replaced every two weeks in both adults and juveniles. The speed of moult in juveniles was generally lower than in adults, in particular of secondaries and tail feathers. Moulting rate of both primaries and tail feathers was lowest in 1994/95 during a period of drought and coincided with the almost complete destruction of roosting habitat. In 1992/93, moulting rate was highest when rainfall was moderate and roosting habitat abundant. Moulting rate was intermediate in 1993/94 when rainfall was frequent but roosting habitat reduced because of the low water level in the Boteti River. The combined effect of reduced food availability during droughts and higher densities and longer foraging flights when roosting habitat is scarce might explain the annual variation in moulting rate. From the second week of January onwards many adults started moulting the outermost tail feather before the penultimate feathers. This phenomenon could indicate the importance of long tail streamers in aerial manoeuvring when foraging during the return migration to the breeding grounds.     

Population response to environmental productivity throughout the annual cycle in a migratory songbird

Environmental factors affect migratory animal populations in every phase of their annual cycle and have significant impacts on breeding success and survival. The Breeding Bird Survey provides a long-term database for examining population trends in North American birds, allowing us to examine large-scale environmental factors that influence population abundance. We examined plant productivity as measured by normalized difference vegetation index (NDVI) over a 24-year period from 1983–2006 in bird conservation regions (BCRs) that overlapped Bullock's oriole (Icterus bullockii) breeding, moult, and wintering ranges to ask whether plant productivity in 1 year influences population abundance in the subsequent breeding season. Bullock's orioles have a moult-migration strategy, with a stopover moult in the Mexican monsoon region, which necessitates examining each stationary phase of the bird's annual cycle to understand the impacts of environmental factors on population abundance. Our results show increased breeding abundance in three (Great Basin, Coastal California and Shortgrass Prairies) of the six BCRs in which the species breeds following years with high NDVI values. We did not detect a response of breeding abundance to high NDVI values in the previous year in either the moulting region or in their primary over-wintering area in central Mexico. Our results demonstrate that large-scale annual variation in primary productivity on the breeding grounds can have an impact on breeding abundance in the following season, but further studies on migratory connectivity and on ecological mechanisms during the non-breeding seasons are needed to understand why we did not detect an influence of productivity during these periods.     

“THE BIRDS OF SOUTH AFRICA”

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

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

Marine moult migration of the freshwater Scaly‐sided Merganser Mergus squamatus revealed by stable isotopes and geolocators

Scaly‐sided Mergansers Mergus squamatus breed on freshwater rivers in far eastern Russia, Korea and China, wintering in similar habitats in China and Korea, but nothing was known of their moulting habitat. To investigate the moult strategies of this species, we combined wing feather stable isotope ratios (males and females) with geolocator data (nesting females) to establish major habitat types (freshwater, brackish or saltwater) used by both sexes during wing moult. Although most Scaly‐sided Mergansers of both sexes probably moult on freshwater, some males and non‐breeding and failed breeding females appeared to undertake moult migration to brackish and marine waters. Given the previous lack of any surveys of coastal or estuarine waters for this species during the moult period, these findings suggest important survey needs for the effective conservation of the species during the flightless moult period.     

Moult of the giant petrels Macronectes halli and M. giganteus at South Georgia

Moult scores were collected from colour-ringed individuals of known reproductive status of the two species of giant petrel, Macronectes halli and M. giganteus , at Bird Island, South Georgia between 1978–81.
Both species showed a substantial overlap between breeding and wing-moult, unlike most other Southern Ocean seabirds. Males started moult before females and both sexes of M. giganteus moulted at an earlier stage of the breeding cycle than M. halli , which breeds six weeks earlier than its congener.
Changes in moult rate during the breeding season are documented for both species, with Id. halli showing a rapid increase as the chick nears fledging. Male M. giganteus show a notably different pattern to the other three species-sex groups, starting moult much earlier (at egg-laying), with greater individual synchrony and usually suspending primary moult throughout the main chick growth period, whereas only two male M. halli and no females of either species suspended moult. Differences in pattern, timing and rate of moult are interpreted in terms of availability of food resources and the competing energy demands of other activities, especially chick-rearing.
Completion of primary moult could not be observed in the field but was estimated using data frcsm non-breeding birds and failed breeders; the latter started a rapid moult almost immediately they failed. In both sexes of both species moult is probably concluded at least by early winter.
The general pattern of moult in giant petrels at Bird Island is contrasted with that of other populations and species of Southern Ocean seabirds. It is suggested that the unusually extensive overlap between breeding and moult in giant petrels is a consequence of the very abundant and easily available summer food supplies (especially carrion) and the much diminished winter resources, favouring a completion of moult by the beginning of the winter.     

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.     

Tracking ice phenology by migratory waterbirds: settling phenology and breeding success of species with divergent population trends

Dependence on climate‐driven environmental cues in the initiation of life cycle stages is a critical attribute when assessing vulnerability of species to climate change impacts. This study focused on spring ice phenology as a cue to the settling of migratory waterbirds, asking whether there is an asynchrony between ice phenology and settling phenology that could affect breeding success of six species with divergent population trends. In the 37 study lakes in southeastern Finland, the ice‐out date not only varied considerably between years, but became progressively earlier during the study period, 1991–2018. Settling phenology of all species tracked inter‐annual variation in ice phenology. However, the degree of asynchrony between ice phenology and settling phenology varied between species, allowing discrimination between early and late settlers. Considerable inter‐annual variation also occurred within species, but in only one species did the degree of asynchrony correlate with the ice‐out date: for the horned grebe Podiceps auritus an earlier ice‐out date meant greater asynchrony between settling phenology and ice phenology. The degree of asynchrony between settling phenology and ice phenology did not affect breeding success in any species. However, ice phenology per se affected breeding success of horned grebes: earlier ice‐out was associated with lower annual breeding success. Breeding numbers of horned grebe showed a long‐term decline. Results suggest that short‐distance migratory birds are able to respond to climate change‐driven phenological changes in their breeding environments, and that this ability may not depend on the relative timing of breeding.     

Feather corticosterone reveals effect of moulting conditions in the autumn on subsequent reproductive output and survival in an Arctic migratory bird

For birds, unpredictable environments during the energetically stressful times of moulting and breeding are expected to have negative fitness effects. Detecting those effects however, might be difficult if individuals modulate their physiology and/or behaviours in ways to minimize short-term fitness costs. Corticosterone in feathers (CORTf) is thought to provide information on total baseline and stress-induced CORT levels at moulting and is an integrated measure of hypothalamic–pituitary–adrenal activity during the time feathers are grown. We predicted that CORTf levels in northern common eider females would relate to subsequent body condition, reproductive success and survival, in a population of eiders nesting in the eastern Canadian Arctic during a capricious period marked by annual avian cholera outbreaks. We collected CORTf data from feathers grown during previous moult in autumn and data on phenology of subsequent reproduction and survival for 242 eider females over 5 years. Using path analyses, we detected a direct relationship between CORTf and arrival date and body condition the following year. CORTf also had negative indirect relationships with both eider reproductive success and survival of eiders during an avian cholera outbreak. This indirect effect was dramatic with a reduction of approximately 30% in subsequent survival of eiders during an avian cholera outbreak when mean CORTf increased by 1 standard deviation. This study highlights the importance of events or processes occurring during moult on subsequent expression of life-history traits and relation to individual fitness, and shows that information from non-destructive sampling of individuals can track carry-over effects across seasons.     

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.     

Assessing the impact of extreme adverse weather on the biological traits of a European storm petrel colony

Climate change affects the climatic disturbance patterns and regimes and is altering the frequency and intensity of subtropical cyclones. These events can affect population dynamics of seabirds (e.g., survival, reproduction). In this work we tested the effect of adverse weather on a colony of European storm petrels (Hydrobates pelagicus) located in a small islet (Aketx) in northern Spain. Over a long-term monitoring period (1993–2014) we ringed 3728 petrels. From 2003 onwards we also monitored breeding success, the percentage of immature individuals and moult scores. We used Cormack-Jolly-Seber models and Underhill and Zucchini models to analyze the effects of climatic conditions on a number of biological traits (survival, breeding parameters, moulting patterns). Our analyses revealed a constant value of adult survival over the 26-year monitoring period. Recapture probability, however, tended to be positively influenced by NAO conditions in spring, and negatively influenced by NAO conditions in winter (although this would only affect to a fraction of first-captured birds). Moreover, the impact of adverse weather, especially in 2011 and 2014, resulted in an increasing proportion of yearlings in the breeding population, a lower breeding success and a delayed onset of moult. These effects were similar to those observed during the Prestige oil spill catastrophe.     

Geographical variation in the timing of breeding and moult in dunlin Calidris alpina on the Palearctic tundra

Studies of how organisms are adapted to regional climatic conditions are valuable when predicting the effects of global climatic changes on biota. Here we report on the geographical variation in timing of breeding and moult of an Arctic breeding wader, the dunlin (Calidris alpina). The Palearctic study sites range latitudinally between 68 and 76°N and longitudinally between 46 and 179°E, and encompass a variety of local climates. The sites were visited in sequence from west to east within 1 year, and therefore the data are not affected by confounding interannual variations. The estimated breeding start ranged from 5 to 25 June across populations. Birds at more southern sites were found to breed earlier than those at more northern breeding sites. Within populations, the breeding start for first clutches spanned a period of 8 days and, when including replacement clutches, 3-4 weeks. No dunlin west of the Taimyr Peninsula were found moulting while incubating at the nest, whereas all dunlin on Taimyr Peninsula and eastwards were in active wing moult while incubating or rearing chicks. The onset of moult in these populations ranged from 23 to 27 June. The consequences of geographical variation of breeding conditions for variation in the annual cycle of this species are discussed.     

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.     

Age and sex differences in the timing of moult in the common seal, Phoca vitulina

This study followed the progress of the annual moult within a population of common seals in Orkney, Scotland. Moulting seals were seen over a three-month period, from 7 June until 16 September. Yearlings were first to start moulting. Amongst older seals, females completed their moult an average of seven days earlier than immature males and 19 days earlier than mature males. Differences in the timing of moult appeared to be related to the age or reproductive status of the animals, and may be the result of differential changes in levels of the sex hormones.     

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.