Energetic consequences of contrasting winter migratory strategies in a sympatric Arctic seabird duet

At the onset of winter, warm‐blooded animals inhabiting seasonal environments may remain resident and face poorer climatic conditions, or migrate towards more favourable habitats. While the origins and evolution of migratory choices have been extensively studied, their consequences on avian energy balance and winter survival are poorly understood, especially in species difficult to observe such as seabirds. Using miniaturized geolocators, time‐depth recorders and a mechanistic model, we investigated the migratory strategies, the activity levels and the energy expenditure of the closely‐related, sympatrically breeding Brünnich's guillemots Uria lomvia and common guillemots Uria aalge from Bjørnøya, Svalbard. The two guillemot species from this region present contrasting migratory strategies and wintering quarters: Brünnich's guillemots migrate across the North Atlantic to overwinter off southeast Greenland and Faroe Islands, while common guillemots remain resident in the Barents, the Norwegian and the White Seas. Results show that both species display a marked behavioural plasticity to respond to environmental constraint, notably modulating their foraging effort and diving behaviour. Nevertheless, we provide evidence that the migratory strategy adopted by guillemots can have important consequences for their energy balance. Overall energy expenditure estimated for the non‐breeding season is relatively similar between both species, suggesting that both southward migration and high‐arctic winter residency are energetically equivalent and suitable strategies. However, we also demonstrate that the migratory strategy adopted by Brünnich's guillemots allows them to have reduced daily energy expenditures during the challenging winter period. We therefore speculate that ‘resident’ common guillemots are more vulnerable than ‘migrating’ Brünnich's guillemots to harsh winter environmental conditions.     

Diving behaviour of guillemot Uria aalge, puffin Fratercula arctica and razorbill Alca tor da as shown by radio-telemetry

Information on dive and pause times and the numbers of dives in a sequence were obtained for six guillemots and single razorbill and puffin. There were marked differences in diving performance between the species with the order of ranking, in descending order of dive and pause duration, being guillemot, razorbill and puffin. For guillemots, 80% of dives were of 20–119 sec duration and 80% of pauses were 0–59 sec; the maximum dive lasted 202 sec. Puffin dives and pauses were much shorter, with 81% of dives lasting 0–39 sec and 95% of pauses being less than 20 sec, the longest dive was 115 sec. Comparisons of diving sequences made by the same individual indicated some flexibility in all aspects of the sequence but there were broad interspecific differences in the organization of the sequence. The puffin generally made a large number of relatively short dives separated by very short pauses which resulted in a high diving rate (1–5 dives/min) and the bird spending 78% of its time underwater. In contrast, guillemots had much shorter sequences with a few long dives and pauses and lower rates of diving (0–5-0-6 dives/min) and proportions of time underwater (61–65%). Guillemots and puffins may forage at different depths and have different foraging strategies.     

Why renew fresh feathers? Advantages and conditions for the evolution of complete post‐juvenile moult

Juveniles of several passerine species renew all of their fresh juvenile feathers immediately after fledging (complete post‐juvenile moult), in contrast to the majority, which perform a partial post‐juvenile moult. To understand the adaptive roles of this phenomenon we compared the quality of juvenile plumage in species that perform a complete post‐juvenile moult with that of species which perform a partial post‐juvenile moult; we similarly compared juveniles and adults in each of these groups. The quality of feathers was measured by mass of primaries, colour, and length. In species which perform a complete post‐juvenile moult the plumage quality of second‐year individuals, in their first breeding season, is similar to the plumage quality of adults, unlike those species that perform a partial post‐juvenile moult. In species which perform complete post‐juvenile moult, the quality of the feathers grown in the nest is lower than the quality of adult post‐breeding feathers. In contrast, in species which perform partial post‐juvenile moult the quality of the feathers grown in the nest is similar to that of adult post‐breeding feathers. We found that a complete post‐juvenile moult strategy is much more common 1) in residents and short‐distance migrants than in long‐distance migrants, 2) in southern latitudes, 3) in species with medium body mass and 4) in omnivores and granivores. Our results indicate two adaptive roles of the complete post‐juvenile moult strategy: 1) achieving high quality plumage in the first year which may increase individual survival probability and fitness and 2) allocating fewer resources to nestling plumage and more to nestling development, which enables the nestlings to leave the nest earlier, thus reducing the probability of encountering nest predators. We suggest that the complete post‐juvenile moult, immediately after fledging, is an optimal strategy in favourable habitats and under low time constraints, as in some tropical ecosystems.     

The stress hormone corticosterone in a marine top predator reflects short‐term changes in food availability

In many seabird studies, single annual proxies of prey abundance have been used to explain variability in breeding performance, but much more important is probably the timing of prey availability relative to the breeding season when energy demand is at a maximum. Until now, intraseasonal variation in prey availability has been difficult to quantify in seabirds. Using a state‐of‐the‐art ocean drift model of larval cod Gadus morhua, an important constituent of the diet of common guillemots Uria aalge in the southwestern Barents Sea, we were able to show clear, short‐term correlations between food availability and measurements of the stress hormone corticosterone (CORT) in parental guillemots over a 3‐year period (2009–2011). The model allowed the extraction of abundance and size of cod larvae with very high spatial (4 km) and temporal resolutions (1 day) and showed that cod larvae from adjacent northern spawning grounds in Norway were always available near the guillemot breeding colony while those from more distant southerly spawning grounds were less frequent, but larger. The latter arrived in waves whose magnitude and timing, and thus overlap with the guillemot breeding season, varied between years. CORT levels in adult guillemots were lower in birds caught after a week with high frequencies of southern cod larvae. This pattern was restricted to the two years (2009 and 2010) in which southern larvae arrived before the end of the guillemot breeding season. Any such pattern was masked in 2011 by already exceptionally high numbers of cod larvae in the region throughout chick‐rearing period. The findings suggest that CORT levels in breeding birds increase when the arrival of southern sizable larvae does not match the period of peak energy requirements during breeding.     

Effects of back-mounted biologgers on condition,diving and flight performance in a breeding seabird

Biologging devices are providing detailed insights into the behaviour and movement of animals in their natural environments. It is usually assumed that this method of gathering data does not impact on the behaviour observed. However, potential negative effects on birds have rarely been investigated before field-based studies are initiated. Seabirds which both fly and use pursuit diving may be particularly sensitive to increases in drag and load resulting from carrying biologging devices. We studied chick-rearing adult common guillemots Uria aalge equipped with and without back-mounted GPS tags over short deployments of a few days. Concurrently guillemots carried small leg-mounted TDR devices (time-depth recorders) providing activity data throughout. Changes in body mass and breeding success were followed for device equipped and control guillemots. At the colony level guillemots lost body mass throughout the chick-rearing period. When-equipped with the additional GPS tag, the guillemots lost mass at close to twice the rate they did when equipped with only the smaller leg-mounted TDR device. The elevated mass loss suggests an impact on energy expenditure or foraging performance. When equipped with GPS tags diving performance, time-activity budgets and daily patterns of activity were unchanged, yet dive depth distributions differed. We review studies of tag-effects in guillemots Uria sp. finding elevated mass loss and reduced chick-provisioning to be the most commonly observed effects. Less information is available for behavioural measures, and results vary between studies. In general, small tags deployed over several days appear to have small or no measurable effect on the behavioural variables commonly observed in most guillemot tagging studies. However, there may still be impacts on fitness via physiological effects and/or reduced chick-provisioning, while more detailed measures of behaviour (e.g. using accelerometery) may reveal effects on diving and flight performance.     

Taking movement data to new depths: Inferring prey availability and patch profitability from seabird foraging behavior

Detailed information acquired using tracking technology has the potential to provide accurate pictures of the types of movements and behaviors performed by animals. To date, such data have not been widely exploited to provide inferred information about the foraging habitat. We collected data using multiple sensors (GPS, time depth recorders, and accelerometers) from two species of diving seabirds, razorbills (Alca torda, N = 5, from Fair Isle, UK) and common guillemots (Uria aalge, N = 2 from Fair Isle and N = 2 from Colonsay, UK). We used a clustering algorithm to identify pursuit and catching events and the time spent pursuing and catching underwater, which we then used as indicators for inferring prey encounters throughout the water column and responses to changes in prey availability of the areas visited at two levels: individual dives and groups of dives. For each individual dive (N = 661 for guillemots, 6214 for razorbills), we modeled the number of pursuit and catching events, in relation to dive depth, duration, and type of dive performed (benthic vs. pelagic). For groups of dives (N = 58 for guillemots, 156 for razorbills), we modeled the total time spent pursuing and catching in relation to time spent underwater. Razorbills performed only pelagic dives, most likely exploiting prey available at shallow depths as indicated by the vertical distribution of pursuit and catching events. In contrast, guillemots were more flexible in their behavior, switching between benthic and pelagic dives. Capture attempt rates indicated that they were exploiting deep prey aggregations. The study highlights how novel analysis of movement data can give new insights into how animals exploit food patches, offering a unique opportunity to comprehend the behavioral ecology behind different movement patterns and understand how animals might respond to changes in prey distributions.     

Ross seal (<Emphasis Type="Italic">Ommatophoca rossii</Emphasis>) annual distribution,diving behaviour,breeding and moulting,off Queen Maud Land,Antarctica

Six out of ten adult Ross seals that were tagged with Argos satellite-linked dive recorders off Queen Maud Land, just after the moult in February, provided data on location and diving activity throughout a year. Shortly after tagging, the animals headed 2,000 km north and stayed pelagic in the area south of the Antarctic Polar Front, until October when they went south into the pack-ice. Throughout the year they made about 100 dives a day, most to a depth of 100–300 m, the deepest dive on record being 792 m, while some dives were very shallow during their stay in the pack-ice. Most dives, outside the breeding and moulting period, lasted for 5–15 min throughout the year. This diving behaviour is consistent with feeding on mid-water fish, like Pleurogramma antarcticum, squid, and to some extent krill (Euphausia superba), when in the pack-ice, and myctophid fish and several species of squid, when in the open ocean. The nursing period was 13 days in mid-November, and moulting occurs in late January–early February, which is the period when sightings surveys for this species should be done.     

Diving behaviour of white-chinned petrels and its relevance for mitigating longline bycatch

The white-chinned petrel (Procellaria aequinoctialis) is the seabird species most commonly killed by Southern Hemisphere longline fisheries. Despite the importance of diving ability for mitigating longline bycatch, little is known of this species’ diving behaviour. We obtained data from temperature–depth recorders from nine white-chinned petrels breeding on Marion Island, southwestern Indian Ocean, during the late incubation and chick-rearing period. Maximum dive depth (16 m) was slightly deeper than the previous estimate (13 m), but varied considerably among individuals (range 2–16 m). Males dived deeper than females, and birds feeding chicks dived deeper than incubating birds, but dive rate did not differ between the sexes. Time of day had no significant effect on dive depth or rate. Our findings will help to improve the design and performance of mitigation measures aimed at reducing seabird bycatch in longline fisheries, such as the calculation of minimum line sink rates and optimum aerial coverage of bird-scaring lines.     

Modelling the Effects of Prey Size and Distribution on Prey Capture Rates of Two Sympatric Marine Predators

Understanding how prey capture rates are influenced by feeding ecology and environmental conditions is fundamental to assessing anthropogenic impacts on marine higher predators. We compared how prey capture rates varied in relation to prey size, prey patch distribution and prey density for two species of alcid, common guillemot (Uria aalge) and razorbill (Alca torda) during the chick-rearing period. We developed a Monte Carlo approach parameterised with foraging behaviour from bird-borne data loggers, observations of prey fed to chicks, and adult diet from water-offloading, to construct a bio-energetics model. Our primary goal was to estimate prey capture rates, and a secondary aim was to test responses to a set of biologically plausible environmental scenarios. Estimated prey capture rates were 1.5±0.8 items per dive (0.8±0.4 and 1.1±0.6 items per minute foraging and underwater, respectively) for guillemots and 3.7±2.4 items per dive (4.9±3.1 and 7.3±4.0 items per minute foraging and underwater, respectively) for razorbills. Based on species'' ecology, diet and flight costs, we predicted that razorbills would be more sensitive to decreases in 0-group sandeel (Ammodytes marinus) length (prediction 1), but guillemots would be more sensitive to prey patches that were more widely spaced (prediction 2), and lower in prey density (prediction 3). Estimated prey capture rates increased non-linearly as 0-group sandeel length declined, with the slope being steeper in razorbills, supporting prediction 1. When prey patches were more dispersed, estimated daily energy expenditure increased by a factor of 3.0 for guillemots and 2.3 for razorbills, suggesting guillemots were more sensitive to patchier prey, supporting prediction 2. However, both species responded similarly to reduced prey density (guillemot expenditure increased by 1.7; razorbill by 1.6), thus not supporting prediction 3. This bio-energetics approach complements other foraging models in predicting likely impacts of environmental change on marine higher predators dependent on species-specific foraging ecologies.     

The post‐fledging period in a tropical bird: patterns of parental care and survival

How environmental conditions affect the timing and extent of parental care is a fundamental question in comparative studies of life histories. The post‐fledging period is deemed critical for offspring fitness, yet few studies have examined this period, particularly in tropical birds. Tropical birds are predicted to have extended parental care during the post‐fledging period and this period may be key to understanding geographic variation in avian reproductive strategies. We studied a neotropical passerine, the western slaty‐antshrike Thamnophilus atrinucha, and predicted greater care and higher survival during the post‐fledging period compared to earlier stages. Furthermore, we predicted that duration of post‐fledging parental care and survival would be at the upper end of the distribution for Northern Hemisphere passerines. Correspondingly, we observed that provisioning continued for 6–12 weeks after fledging. In addition, provisioning rate was greater after fledging and offspring survival from fledging to independence was 75%, greater than all estimates from north‐temperate passerines. Intervals between nesting attempts were longer when the first brood produced successful fledglings compared to nests where offspring died either in the nest or upon fledging. Parents delayed initiating second nests after the first successful brood until fledglings were near independence. Our results indicate that parents provide greater care after fledging and this extended care likely increased offspring survival. Moreover, our findings of extended post‐fledging parental care and higher post‐fledging survival compared to Northern Hemisphere species have implications for understanding latitudinal variation in reproductive effort and parental investment strategies.     

Inter-breeding movements of common guillemots (Uria aalge) suggest the Barents Sea is an important autumn staging and wintering area

Seabird movements outside the breeding season are generally poorly known, but can cover thousands of square km and a multitude of habitats, feeding conditions and potential threats. During the last decades, many seabird species in the North Atlantic have experienced large reductions in population size and breeding success, probably caused by reduced prey abundance caused by climate alterations and overfishing. One of these seabird species is the common guillemot. We used global location sensors (geolocators) to identify inter-breeding movements of 10 individuals breeding at Sklinna, a colony off the coast of Central Norway during July 2009–July 2010. All individuals moved northwards after breeding, and eight of them (80?%) entered the Barents Sea where they probably completed their moult. Three individuals moved southwards before the winter, but in total, half of the individuals stayed in the Barents Sea during winter. The other half wintered off the coast of Central Norway–Lofoten. The fact that all individuals moved northwards to winter was surprising as ringing recoveries suggest they also moves southwards (to the Skagerrak area) to winter. This suggests variation (individual or annual) in wintering movements and calls for a multi-year geolocator study at a number of colonies. Much of the area in the Barents Sea–Lofoten area is classified as vulnerable with respect to specific environmental pressures such as oil pollution and other anthropogenic factors, and the importance of the Barents Sea as a major wintering area for common guillemots from central Norway certainly has implications for the management authorities.     

The diving behaviour of the Manx Shearwater Puffinus puffinus

The diving capabilities of the Procellariformes remain the least understood component of avian diving physiology. Due to their relatively small size, shearwaters may have high oxygen consumption rates during diving relative to their available oxygen stores. Dive performance in this group should be strongly limited by the trade‐off between oxygen consumption and oxygen stores, and shearwaters could be a good model group for testing predictions of dive theory. Many earlier measurements of shearwater dive behaviour relied on observations from the surface or potentially biased technology, and it is only recently that diving behaviour has been observed using electronic recorders for many of the clades within the family. The diving behaviour of Manx Shearwaters Puffinus puffinus breeding in Wales, UK, was studied on a large sample of birds using time–depth–temperature recorders deployed on chick‐rearing shearwaters in July and August over 3 years (2009–2011). Light availability apparently limited diving as dives only occurred between 04:00 and 19:00 h GMT. All individuals routinely dived deeper than traditionally assumed, to a mean maximum depth of 31 m and occasionally down to nearly 55 m. We compiled all available data for a comparison of the dive depth across shearwater species. There was a positive allometric relationship between maximum dive depth and body mass across Puffinus and Ardenna shearwater species, as expected, but only if samples of fewer than two individuals were excluded. The large intra‐specific range in maximum dive depth in our study illustrates that apparent diversity in diving performance across species must be interpreted cautiously.     

Post‐fledging survival of altricial birds: ecological determinants and adaptation

For altricial young, fledging is an abrupt step into an unknown environment. Despite increasing numbers of studies addressing the post‐fledging period, our current knowledge of the causes and consequences of post‐fledging survival remains fragmentary. Here, we review the literature on post‐fledging survival of juvenile altricial birds, addressing the following main questions: Is low post‐fledging survival a bottleneck in the altricial reproductive cycle? What is known of proximate and ultimate causal factors such as trophic relations (food and predation), habitat conditions, or abiotic factors acting in the post‐fledging period? We analyzed weekly survival estimates from 123 data series based on studies of 65 species, covering weeks 1–13 post‐fledging. As a general pattern, survival of fledglings was low during the first week post‐fledging (median rate = 0.83), and improved rapidly with time post‐fledging (week 4 median rate = 0.96). For ground‐nesting species, survival immediately after leaving nests was similar to egg‐to‐fledging survival. For species breeding above‐ground, survival during the first week post‐fledging was substantially lower than during both the nestling period and later post‐fledging stages. Thus, the early post‐fledging period is a bottleneck of markedly elevated mortality for most altricial species. Predation was the main proximate cause of mortality. Various factors such as habitat, annual and seasonal variation in the environment, and the physical condition of fledglings have been found to affect post‐fledging survival. Individual survival depended strongly on physical traits such as mass and wing length, which likely influence the ability of fledglings to escape predation. Trophic relationships at various levels are the main ultimate driver of adaptation of traits relevant to survival during the pre‐ and post‐fledging periods. Spatiotemporal dynamics of food resources determine the physical development of juveniles and, in turn, their performance after fledging. However, predators can cause quick and efficient selection for fledgling traits and adult breeding decisions. Parental strategies related to clutch size and timing of breeding, and the age and developmental stage at which young fledge have substantial effects on post‐fledging survival. The intensity and duration of post‐fledging parental investment also influences fledgling survival. Post‐fledging mortality is therefore not a random and inevitable loss. Traits and strategies related to fledging and the post‐fledging stage create large fitness differentials and, therefore, are integral, yet poorly understood, parts of the altricial reproductive strategy.     

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.     

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

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

Species‐specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle

Determining the year‐round distribution and behaviour of birds is necessary for a better understanding of their ecology and foraging strategies. Petrels form an important component of the high‐latitude seabird assemblages in terms of species and individuals. The distribution and foraging ecology of three sympatric fulmarine petrels (Southern Fulmar Fulmarus glacialoides, Cape Petrel Daption capense and Snow Petrel Pagodroma nivea) were studied at Adélie Land, East Antarctica, by combining information from miniaturized saltwater immersion geolocators and stable isotopes from feathers. During the breeding season at a large spatial scale (c. 200 km), the three species overlapped in their foraging areas located in the vicinity of the colonies but were segregated by their diet and trophic level, as indicated by the different chick δ¹⁵N values that increased in the order Cape Petrel < Southern Fulmar < Snow Petrel. During the non‐breeding season, the three fulmarines showed species‐specific migration strategies along a wide latitudinal gradient. Snow Petrels largely remained in ice‐associated Antarctic waters, Southern Fulmars targeted primarily the sub‐Antarctic zone and Cape Petrels migrated further north. Overall, birds spent less time in flight during the non‐breeding period than during the breeding season, with the highest percentage of time spent sitting on the water occurring during the breeding season and at the beginning of the non‐breeding period before migration. This activity pattern, together with the δ¹³C values of most feathers, strongly suggests that moult of the three fulmarine petrels occurred at that time in the very productive high Antarctic waters, where birds fed on a combination of crustaceans and fish. The study highlights different segregating mechanisms that allow the coexistence of closely related species, specifically, prey partitioning during the breeding season and spatial segregation at sea during the non‐breeding season.     

Late‐moulting Black‐necked Grebes Podiceps nigricollis show greater body mass in the face of failing food supply

From August to December, thousands of Black‐necked Grebes Podiceps nigricollis concentrate during the flightless moult period in salt ponds in the Odiel Marshes, southern Spain, where they feed on the brine shrimp Artemia parthenogenetica. We predicted that because Black‐necked Grebes moulted in a food‐rich, predator‐free environment, there would be no net loss of body mass caused by the use of fat stored to meet energy needs during remigial feather replacement (as is the case for some other diving waterbirds). However, because the food resource disappears in winter, we predicted that grebes moulting later in the season would put on more body mass prior to moult because of the increasing risk of an Artemia population crash before the moult period is completed. Body mass determinations of thousands of birds captured during 2000–2010 showed that grebes in active wing‐moult showed greater mass with date of capture. Early‐moulting grebes were significantly lighter at all stages than late‐moulting birds. Grebes captured with new feathers post‐moult were significantly lighter than those in moult. This is the first study to support the hypothesis that individual waterbirds adopt different strategies in body mass accumulation according to timing of moult: early‐season grebes were able to acquire an excess of energy over expenditure and accumulate fat stores while moulting. Delayed moulters acquired greater fat stores in advance of moult to contribute to energy expenditure for feather replacement and retained extra stores later, most likely as a bet hedge against the increasing probability of failing food supply and higher thermoregulatory demands late in the season. An alternative hypothesis, that mass change is affected by a trophically transmitted cestode using brine shrimps as an intermediate host and Black‐necked Grebes as final host, was not supported by the data.     

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

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

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.