Seasonal variation in food availability influences the breeding strategy of White‐collared Blackbirds Turdus albocinctus on the Tibetan Plateau

Parent birds show a continuous spectrum of breeding strategies, ranging from a low‐fecundity and high‐survival pattern to a high‐fecundity, low‐survival pattern. Investigations of parental breeding strategies under variable environmental conditions can illustrate how parents trade‐off the benefits and costs of these two extreme strategies. White‐collared Blackbirds Turdus albocinctus can breed twice a year on the Tibetan Plateau. We show that both life‐history traits and parental feeding behaviour differ between these two breeding attempts. In the first attempt, the birds produced small clutches and fledged a small number of nestlings of high body condition. In the second attempt, they produced larger clutches and fledged more nestlings of lower body condition. Males made greater contributions to brood provisioning compared with females in the first attempt but there was no sex difference in brood provisioning in the second attempt. In the first attempt, producing smaller clutches can shorten the nestling period, and the increased male contribution to brood provisioning can protect the energy reserves of females. Thus, females can begin a second attempt sooner and produce larger clutches. During the second nesting attempt, when conditions are warmer and wetter, parents rely on a broader array of food types (both invertebrates and plant material, primarily berries) than during the first attempt, which includes only animal food such as arthropods and annelids. We suggest that this difference in breeding strategies between nesting attempts and sexes is in part influenced by marked seasonal variation in food availability.     

Brood parasitism by cowbirds: risks and effects on reproductive success and survival in indigo buntings

We observed brood parasitism by brown-beaded cowbirds (Molothrusater) on indigo buntings (Passerina cyanea) and estimated dieimpact of parasitism on the success of the individual buntingsin their current nests and in their future survival and reproduction.Rates of parasitism over 8 years were 26.6% in 1040 nests and19.8% in 693 nests in two areas in southern Michigan. Risk ofparasitism was high early in the season; half the bunting nestswere begun after the end of the cowbird season. Risk was independentof female age, plant containing the nest, or habitat The immediatecost of parasitism was 1.19 and 1.26 fewer buntings fledgedper nest. Bunting success was lower in parasitized nests withcowbird eggs (nests were more likely to be deserted or predated),lower when the cowbird nestling failed (nests were more likelyto be predated), and lower when the cowbird fledged (fewer buntingsfledged) compared to nonparasitized nests. Costs were due toremoval of a bunting egg when die cowbird laid its own egg andto competition for parental care of the cowbird and buntingnestlings. Buntings that fledged from nests where a cowbirdalso fledged were only 18% as likely to survive and return totheir natal area in the next year as buntings from nests wherea cowbird did not fledge. Long-term effects of cowbird parasitismon adult breeding later in the season, survival to the nextseason, and reproductive success in the next season were negligiblewhen compared between birds that reared a cowbird and birdsthat reared only a bunting brood, or between birds that wereparasitized and birds that escaped parasitism. The results indicatelittle long-term cost of brood parasitism on individual fitnessof adult buntings beyond the impact on the current nest andthe survival of buntings that fledge from it; nearly all costis to the parasitized brood.     

The Process and Causes of Fledging in a Cavity-Nesting Passerine Bird, the House Wren (Troglodytes aedon)

Little is known about the process or causes of fledging or nest‐leaving in passerine birds because researchers can rarely predict when fledging will occur in a given nest. We used continuous videotaping of nests to both document the process of fledging in the house wren, Troglodytes aedon, a small, cavity‐nesting songbird, and test hypotheses as to what might cause fledging to begin. Fledging began any time from 14 to 19 d after hatching commenced. Slower‐developing broods fledged later than faster‐developing broods. Fledging typically began within 5 h of sunrise and over 80% of all nestlings fledged before noon. All nestlings fledged on the same day at 65% of nests and over two consecutive days in most other nests. We found no evidence that fledging was triggered by changes in parental behaviour. Parental rate of food delivery to nestlings did not decline during a 3‐h period leading up to the first fledging, nor was the rate of feeding just prior to the first fledging lower than the rate at the same time the day before. Moreover, parents did not slow the rate of food delivery to nests after part of the brood had fledged. Hatching is asynchronous in our study population which creates a marked age/size hierarchy within broods. At most nests, the first nestling to fledge was the most well‐developed nestling in the brood or nearly so (as measured by feather length). This suggests that fledging typically begins when the most well‐developed nestlings in the brood reach some threshold size. However, at about one‐fifth of nests, the first nestling to fledge was only moderate in size. At these nests, severe competition for food may have caused smaller, less competitive nestlings to fledge first to increase their access to food. We found no strong support for the suggestion that the oldest nestlings delay fledging until their least‐developed nestmate reaches some minimum size, although further experimental work on this question is warranted.     

The presence of kleptoparasitic fledglings is associated with a reduced breeding success in the host family in the barn owl

Fledgling birds sometimes abandon their own nest and move to neighboring nests where they are fed by host parents. This behaviour, referred to as ‘nest‐switching’, is well known in precocial birds that are mobile soon after hatching and can easily reach foster nests. In contrast, due to the difficulty of observing nest‐switching in territorial altricial birds, the causes and consequences of moving to others’ nests are poorly known in this group of birds. Nest‐switchers can be adopted by the foster parents or they can steal food from the host parents meant for their offspring, a form of kleptoparasitism, which may result in reduced breeding success of the host nest. In Israel, 12 barn owl fledglings left their natal nests and were found in 9 host nests out of 111 monitored nests (8.1%). Nest‐switchers that fledged earlier in the breeding season flew shorter distances to reach host nests probably because the density of nests with younger nestlings is higher early in the season. The number of host nestlings fledged and the percentage of nestlings fledged was lower in host nests than in nests without switchers. The occasional nest‐switchers were always older than host nestlings (respectively 80 and 50 days of age, on average) and host parents fledged fewer young when nest‐switchers occupied host nests with younger nestlings. This suggests that nest‐switchers are kleptoparasites because the presence of the older alien fledglings is associated with a lower breeding success of the host parents.     

Effects of geolocator attachments on breeding parameters of Lesser Kestrels

ABSTRACT Light level geolocators, also known as GLS loggers, are electronic devices intended for tracking the location of wide‐ranging animals using ambient light to estimate latitude and longitude. Miniaturized geolocators that can be used on relatively small migratory birds have recently become available, but little is known about the potential harmful effects of geolocators on birds. We examined the possible effects of 1.5‐g geolocators (dimensions: 21 × 6.5 × 9 mm) on the breeding success and survival of migratory Lesser Kestrels (Falco naumanni). During the 2007 breeding season, kestrels were fitted with geolocators using two attachment methods (Teflon wing harnesses and darvic bands), and geolocators were removed in 2008 after the birds returned to the breeding grounds. We found no differences in the breeding success of control and tagged pairs during the 2007 breeding season, but tagged pairs had greater fledgling mortality in the following breeding season. Furthermore, nestlings of tagged individuals had higher triglyceride and uric acid concentrations in their blood than control nestlings during the breeding season following tagging. As for return rates, 75% of tagged birds came back to the colony after the nonbreeding period, a proportion similar to that reported in previous studies. Although back‐mounts are slightly heavier and require more skill to attach, we recommend their use on small migratory raptors because most leg‐mounted geolocators in this study were damaged or rendered useless by dirt obscuring the light sensor.     

Parasites favour intermediate nestling mass and brood size in cliff swallows

A challenge of life‐history theory is to explain why animal body size does not continue to increase, given various advantages of larger size. In birds, body size of nestlings and the number of nestlings produced (brood size) have occasionally been shown to be constrained by higher predation on larger nestlings and those from larger broods. Parasites also are known to have strong effects on life‐history traits in birds, but whether parasitism can be a driver for stabilizing selection on nestling body size or brood size is unknown. We studied patterns of first‐year survival in cliff swallows (Petrochelidon pyrrhonota) in western Nebraska in relation to brood size and nestling body mass in nests under natural conditions and in those in which hematophagous ectoparasites had been removed by fumigation. Birds from parasitized nests showed highest first‐year survival at the most common, intermediate brood‐size and nestling‐mass categories, but cliff swallows from nonparasitized nests had highest survival at the heaviest nestling masses and no relationship with brood size. A survival analysis suggested stabilizing selection on brood size and nestling mass in the presence (but not in the absence) of parasites. Parasites apparently favour intermediate offspring size and number in cliff swallows and produce the observed distributions of these traits, although the mechanisms are unclear. Our results emphasize the importance of parasites in life‐history evolution.     

The effects of force‐fledging and premature fledging on the survival of nestling songbirds

Despite the broad consensus that force‐fledging of nestling songbirds lowers their probability of survival and therefore should be generally avoided by researchers, that presumption has not been tested. We used radiotelemetry to monitor the survival of fledglings of Ovenbirds Seiurus aurocapilla and Golden‐winged Warblers Vermivora chrysoptera that we unintentionally force‐fledged (i.e. nestlings left the nest in response to our research activities at typical fledging age), that fledged prematurely (i.e. nestlings left the nest earlier than typical fledging age), and that fledged independently of our activities. Force‐fledged Ovenbirds experienced significantly higher survival than those that fledged independent of our activities, and prematurely fledged Ovenbirds had a similarly high survival to those that force‐fledged at typical fledging age. We observed a similar, though not statistically significant, pattern in Golden‐winged Warbler fledgling survival. Our results suggest that investigator‐induced force‐fledging of nestlings, even when deemed premature, does not necessarily result in reduced fledgling survival in these species. Instead, our results suggest that a propensity or ability to fledge in response to disturbance may be a predictor of a higher probability of fledgling survival.     

Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

An experimental approach to the brood reduction hypothesis in Magellanic penguins

In many bird species, eggs in a brood hatch within days of each other, leading to a size asymmetry detrimental to younger siblings. Hatching asynchrony is often thought of as an adaptive strategy, and the most widely studied hypothesis in relation to this is the ‘brood reduction hypothesis’. This hypothesis states that when food resources are unpredictable, hatching asynchrony will allow the adjustment of the brood size maximizing fledging success and benefitting parents. The Magellanic penguin Spheniscus magellanicus is an appropriate species to test this hypothesis because it has a 2‐egg clutch that hatches over a 2‐d interval with a broad range of variation (–1 to 4 d), it shows facultative brood reduction, and food abundance between breeding seasons is variable. We performed a manipulative study at Isla Quiroga, Argentina, during three breeding seasons (2010–2012) by forcing broods to hatch synchronously (0 d) or asynchronously (2 or 4 d). Years were categorized based on estimated food abundance. Our study provided mixed results because in the low estimated food abundance year asynchronous broods did not have higher nestling survival than synchronous broods, and the second‐hatchling in asynchronous broods did not die more often than those in synchronous broods. On the other hand, younger siblings of 4‐d asynchronous broods starved earlier than those of synchronous broods, and 2‐d asynchronous broods fledged heavier young than synchronous broods. Asynchronous hatching would seem to benefit reproduction in this species, not with respect to survival, but in terms of the advantages it can accord to nestlings and, in terms of lower costs, for parents raising nestlings.     

Dispersal and recruitment during population growth in a colonial bird, the great cormorant Phalacrocorax carbo sinensis

While the factors influencing reproduction and survival in colonial populations are relatively well studied, factors involved in dispersal and settlement decisions are not well understood. The present study investigated exchanges of great cormorants Phalacrocorax carbo sinensis among six breeding colonies over a 13‐year period when the breeding population in Denmark increased from 2800 to 36 400 nests. We used a multistate capture‐recapture model that combined multisite resightings and recoveries to examine simultaneously recruitment, natal dispersal, breeding dispersal and annual survival of first‐year, immature and breeding great cormorants. Mean survival of first‐year birds (0.50±0.09, range=0.42–0.66 among colonies) was lower than survival of breeders (0.90±0.06, range=0.81–0.97). Mean survival of immature birds over the study period was 0.87±0.08. Dispersal from a colony increased with decreasing mean brood size in the colony in both first‐time and experienced breeders. The choice of the settlement colony in first‐time breeders was affected by conditions in the natal colony and in the colonies prospected during the pre‐breeding years. In particular, first‐time breeders recruited to colonies where they could expect better breeding success. Experienced breeders relied mainly on cues present early in the season and on their own breeding experience to choose a new breeding colony. Newly established colonies resulted mainly from the immigration of first‐time breeders originating from denser colonies. Dispersal was distance‐dependent and first‐time breeders dispersed longer distances than breeders. We suggest that the prospecting behaviour allows first‐time breeders to recruit in nearby as well as more distant potential breeding colonies. Dispersing breeders preferred to settle in neighbouring colonies likely to benefit from their experience with foraging areas. We discuss the importance of these movements for growth and expansion of the breeding population.     

Costs of rearing and sex‐ratio variation in southern grey shrike Lanius meridionalis broods

Several non‐mutually exclusive hypotheses predict adaptive variation in the offspring sex ratio. When conditions for breeding are adverse, parents are predicted to produce more offspring of the less costly sex to rear (‘the cost‐of‐reproduction hypothesis’). Moreover, they also should produce the more dispersing sex in order to diminish future competition (‘the local‐resource‐competition hypothesis’). Here, we analyse brood sex ratio according to rearing conditions in the southern shrike Lanius meridionalis, a species with moderately reversed sexual dimorphism. Our results suggest that females are more costly to rear than males in this species. Adult females proved heavier than males, and female nestling tended to be heavier than male nestlings. Moreover, the greater brood reduction, the more male‐biased was the brood, suggesting that brood reduction implied higher mortality in female nestlings. Consistent with these findings, the brood sex ratio was biased to the less costly sex (males) when breeding conditions were adverse (bad years or low‐quality male parents), supporting the cost‐of‐reproduction hypothesis. By contrast, these findings did not support the local‐resource‐competition hypothesis, which predicted female‐biased brood sex ratio under adverse conditions. As a whole, our results support the idea that birds adaptively modulate sex ratio in order to minimize reproduction costs.     

Behaviour and breeding biology of the cooperatively breeding Grey-backed Fiscal Shrike Lanius excubitorius in Kenya

Grey-backed Fiscal Shrikes Lanius excubitorius were studied over a 2j year period near Lake Naivasha, Kenya. Grey-backs are cooperative breeders, with group sizes ranging from two to II. Only one pair breeds per group, with all other group members aiding in the rearing of young. The study population ranged from 64 to 79 individuals that occurred in from 13 to 16 groups. Non-breeding helpers made up to 66% of the population, with male helpers being more numerous overall than females. The annual survival rate was 65%, with no differences detected between the survival of males and females, or of breeders and helpers. Only male helpers were observed to acquire breeding status within the natal territory. Some female helpers acquired breeding positions in territories adjacent to their natal territories. Group territorial displays occurred throughout the year but were most pronounced prior to breeding during rainy periods. Reproductive success was very low, with only 14.5% of the recorded breeding attempts leading to fledged young. Large groups (four or more birds) had greater reproductive success than small groups, but many factors other than, or in addition to, group size may have influenced this pattern. The breeding male contributed the most food to the incubating female and to the nestlings. Male helpers and the breeding female contributed more to nestlings than did female helpers. Observations on the post-fledging period indicate that socialization and establishment of dominance may be of importance in group-living species living in a restricted ecological and social setting.     

Effect of food availability on nestling growth and fledging success in manipulated pallid swift broods

Growth rate and fledging success were assessed in natural and manipulated broods of the pallid swift Apus pallidus. Daily measurements of chick mass, wing length, and insect abundance allowed us to examine the short-term variation of chick growth in relation to food availability.
The number of fledged nestlings increased with brood size. Wing length and body mass were slightly but significantly smaller in larger broods, and the nestlings of enlarged broods needed longer to fledge. We discuss how these differences could influence survival after fledging.
Hatching asynchrony caused a significant difference in growth among siblings, and the difference between the oldest and youngest chick was greater in larger broods.
Chick growth was independent of daily food availability. We suggest that this was due to an increased effort of the parents at their expense, when food availability was poor.
The ability of this species to raise an additional chick is in line with most findings on birds, but partially in contrast with results for the common swift in which, at least during poor seasons, the additional nestling caused an increased mortality and lowered the reproductive success.     

Carry‐over effects provide linkages across the annual cycle of a Neotropical migratory bird,the Louisiana Waterthrush Parkesia motacilla

Population limitation models of migratory birds have sought to include impacts from events across the full annual cycle. Previous work has shown that events occurring in winter result in some individuals transitioning to the breeding grounds earlier or in better physical condition than others, thereby affecting reproductive success (carry‐over effects). However, evidence for carry‐over effects from breeding to wintering grounds has been shown less often. We used feather corticosterone (CORT_f) levels of the migratory Louisiana Waterthrush Parkesia motacilla as a measure of the physiological state of birds at the time of moult on the breeding territory to investigate whether carry‐over effects provide linkages across the annual cycle of this stream‐obligate bird. We show that birds arriving on wintering grounds with lower CORT_f scores, indicating reduced energetic challenges or stressors at the time of moult, occupied higher quality territories, and that these birds then achieved a better body condition during the overwinter period. Body condition, in turn, was important in determining whether adult birds returned the following winter, with birds in better condition returning at higher rates. Together these data suggest a carry‐over effect from the breeding grounds to the wintering grounds that is further extended with respect to annual return rates. Very few other studies have linked conditions during the previous breeding season with latent effects during the subsequent overwintering period or with annual survival. This study shows that the effects of variation in energetic challenges or stressors can potentially carry over from the natal stream and accumulate over more than one life‐history period before being manifested in reduced survival. This is of particular relevance to models of population limitation in migratory birds.     

Age‐related prenatal maternal effects and postnatal breeding experience have different influences on nestling development in an altricial passerine

Reproductive success and nestling performance are related to the age of parents across several vertebrate taxa. However, because breeding experience and prenatal maternal investment in reproduction often covary, the source of these age‐related differences can be difficult to determine. In this study, we evaluated the influence of prenatal maternal effects and postnatal breeding experience on the performance of nestling tree swallows Tachycineta bicolor by conducting a carefully controlled partial cross‐fostering experiment. We swapped half‐broods of nestlings between the nest of a young first‐time breeding female and the nest of a female known to have previously raised and fledged young. Our manipulation did not influence the within‐brood nestling hierarchies, and controlled for the effects of egg laying order. We found that nestlings of older females were heavier just prior to fledging regardless of the breeding experience of the attending female. In addition, fledglings raised by experienced females grew their flight feathers faster, and had greater probability of fledging. Our study demonstrates that prenatal investment in reproduction by older females can have long‐term consequences on nestling mass, and suggests limited potential for compensatory mass gains prior to fledging. Because our analyses controlled for feeding rates, our results also suggest that foraging quantity and quality are not the only benefits nestlings gain by being raised by an experienced female.     

Hatching asynchrony and growth trade‐offs within domesticated and wild zebra finch,Taeniopygia guttata,broods

The Australian zebra finch, Taeniopygia guttata, is a widely used model organism, yet few studies have compared domesticated and wild birds with the aim of examining its relevance as an evolutionary model species. Domestic and wild broods hatch over approximately 4 and 2 days, respectively, which is important given that nestlings can fledge after as little as 12 days, although 16–18 days is common. We aimed to evaluate the extent to which the greater hatching asynchrony in domestic stock may effect reproductive success through greater variance in size hierarchies, variance in within‐brood growth rates, and partial brood mortality. Therefore, by simultaneously controlling brood sizes and experimentally manipulating hatching intervals in both domesticated and wild birds, we investigated the consequences of hatching intervals for fledging success and nestling growth patterns, as well as trade‐offs. Fledging success was similarly high in domestic and wild broods of either hatching pattern. Nonetheless, between‐brood analyses revealed that domestic nestlings had significantly higher masses, larger skeletal characters, and longer wings than their wild counterparts, although wild nestlings had comparable wing lengths at the pre‐fledging stage. Moreover, within‐brood analyses revealed only negligible differences between domestic and wild nestlings, and larger effects of hatching order and hatching pattern. Therefore, despite significant differences in the hatching intervals, and the ultimate size achieved by nestlings, the domestication process does not appear to have significantly altered nestling growth trade‐offs. The present study provides reassuring evidence that studies involving domesticated zebra finches, or other domesticated model organisms, may provide reasonable adaptive explanations in behavioural and evolutionary ecology. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 763–773.     

Long-term consequences of early ontogeny in free-living Great Tits Parus major

Environmental factors during early development may have profound effects on subsequent life-history traits in many bird species. In wild birds, sex-specific effects of early ontogeny on natal dispersal and future reproduction are not well understood. The objective of this work was to determine whether hatching date and pre-fledging mass and condition of free-living Great Tits Parus major have any subsequent effect on individuals’ natal dispersal and reproductive performance at first breeding. Both males and females dispersed longer distances in coniferous than in deciduous forests, while dispersal was condition-dependent only in males (heavier as nestlings dispersed farther). In females, mass and condition at pre-fledging stage correlated significantly with clutch size, but not with subsequent reproductive performance as measured by fledging success or offspring quality. In contrast, heavier males as nestlings had higher future fledging success and heavier offspring in their broods compared with those in worse condition as nestlings. The hatching date of female as well as male parents was the only parental parameter related to the number of eggs hatched at first breeding. These results indicate that pre-fledging mass and condition predict subsequent fitness components in this bird species. We suggest that sex-specific relationships between a disperser’s condition and its selectivity with respect to breeding habitat and subsequent performance need to be considered in future models of life-history evolution.     

Individual variation in parental tradeoffs between the number and size of offspring at the pre- and post-natal stages

Life-history theory predicts that parents refer to the resources they hold to determine their breeding strategy. In multi-brooded species, it is hypothesized that single-brooded parents produce larger clutches and raise offspring with a brood survival strategy, whereas multi-brooded parents only do this under good breeding conditions. Under poor conditions, they produce smaller clutches and raise offspring with a brood reduction strategy. We tested this hypothesis in the Brown-cheeked Laughing Thrush Trochalopteron henrici, which can breed twice a year on the Tibetan Plateau, by investigating the life-history traits and provisioning behaviours of single- and double-brooded parents. Single-brooded parents laid larger clutches of smaller eggs and produced more and larger fledglings than double-brooded parents in their first brood. Double-brooded parents produced smaller clutches of larger eggs but fledged larger nestlings in their first brood than in their second brood. As single-brooded parents only need to raise one brood a year, then producing and raising as many offspring as possible (i.e. the brood survival strategy in a large brood) can maximize their reproductive success. For double-brooded parents, producing and raising fewer offspring in the first brood (i.e. the brood survival strategy in a small brood) can ensure their nesting success during a short breeding cycle. Additionally, producing more offspring but raising larger nestlings in the second brood (i.e. the brood reduction strategy in a large brood) can select for offspring of higher quality within the brood. Our findings indicate that different tradeoffs between single- and double-brooded parents in egg-laying and nestling-raising may be an adaptation to the seasonal variation in environmental conditions.     

Nest helpers improve parental survival but not offspring production in a high‐elevation passerine,the Ground Tit Pseudopodoces humilis

The way in which breeders respond to helping, in terms of either offspring production or their own survival, may reflect the adaptive aspects of a cooperative breeding system. We explore this issue using a 5‐year study of the Ground Tit Pseudopodoces humilis, a facultative cooperative breeder in which 47% of socially monogamous pairs have between one and four close male relatives as helpers. We found that helped nests did not fledge more or heavier nestlings than unhelped nests, and male young from helped and unhelped nests were equally likely to recruit into the local breeding population. However, helped parents of both sexes had a higher probability of survival to the following year than did unhelped parents. These findings suggest that Ground Tit parents with helpers trade current reproduction for personal survival and future reproduction, a strategy favoured by selection to cope with harsh, unpredictable environments such as the Tibetan Plateau.     

Sex,growth rate,rank order after brood reduction,and hatching date affect first‐year survival of long‐lived Herring Gulls

Among most species of birds, survival from hatching throughout the first year of life is generally lower than subsequent survival rates. Survival of young birds during their first year may depend on a combination of selection, learning, unpredictable resources, and environmental events (i.e., post‐fledging factors). However, knowledge about post‐fledging development in long‐lived species is usually limited due to a lengthy immature stage when individuals are generally unobservable. Therefore, pre‐fledging characteristics are often used to predict the survival of young birds. We assessed effects of nestling growth rates, hatching date, hatching asynchrony, brood size and rank order after brood reduction, and sex on first‐year survival of 137 fledglings using a mark‐resighting analysis. We found that the survival probability (Φ_1yr = 0.39) of first‐year Herring Gulls (Larus argentatus) in our study colony located at the outer port of Zeebrugge (Belgium) was lower than that of older individuals (Φ_>1yr = 0.75). All 10 models best supported by our data included nestling growth rate, suggesting that variability in first‐year survival may be linked primarily to individual variation in growth. First‐year survival was negatively correlated with hatching date and rank order after brood reduction. Hence, carry‐over effects of breeding season events such as timing of breeding, early development, and social status had an influence on survival of Herring Gulls after fledging. Furthermore, we found sex‐biased mortality in first‐year Herring Gulls, with females (Φ_1yr = 0.45) surviving better than males (Φ_1yr = 0.38). Although adult survival is generally regarded as the key parameter driving population trajectories in long‐lived species, juvenile survival has recently been acknowledged as an important source of variability in population growth rates. Thus, increasing our knowledge of factors affecting age‐specific survival rates is necessary to improve our understanding of population dynamics and ultimately life‐history variation.