Do environmental conditions experienced in early life affect recruitment age and performance at first breeding in common goldeneye females?

Environmental conditions experienced early in life may have long‐term impacts on life history traits and reproductive performance. We investigated whether ambient temperature experienced during the first two to four weeks of life and weather severity during the first two winters affected recruitment age and relative timing of breeding in the year of recruitment in female common goldeneyes Bucephala clangula. Our sample consisted of 141 female recruits hatched in a study population in central Finland between 1985 and 2013 and captured later as breeders. About 56% of the recruited females bred for the first time when two years old (range 2–6 yr). Individuals facing colder ambient temperatures during the first two to four weeks posthatch or more severe winter conditions during the first two winters did not recruit at an older age. Nor did maternal characteristics, relative hatch date or nest site availability affect recruitment age. For females that recruited at two years old, the date of first breeding was usually late relative to the population mean that year (mean difference 6.9 d, range –7 to 21 d). Our results suggest developmental buffering enables female goldeneye ducklings to mitigate the impacts of adverse environmental conditions experienced during the first weeks of life, at least in terms of first breeding.     

Brood rearing costs affect future reproduction in the precocial common goldeneye Bucephala clangula

Life-history theory assumes a trade-off between current reproductive effort and future reproductive success. There are a large number of studies demonstrating reproductive trade-offs in different animal taxa, particularly in birds. Most bird studies have focused on the costs of chick rearing in altricial species. These costs have been assumed to be low in precocial species, but this aspect has been little studied. We used long-term individual reproductive data from the common goldeneye Bucephala clangula , an iteroparous precocial duck with uniparental female care, to examine whether brood rearing carries costs that affect future reproductive performance. All females were experienced breeders, and possible differences in female quality were ruled out. We compared within-individual (between-year) changes in clutch size, hatching date and body mass between females that had reared a brood in the previous year and females that had not. It turned out that brood rearing involved a cost in terms of clutch size and hatching date the next year, but not in terms of body mass: females that had reared a brood in the previous year laid relatively smaller clutches and laid relatively later than females that had not reared a brood. Our results show that normal brood rearing in a precocial species involves costs that affect future reproduction.     

Landscape-level correlates of mallard duckling survival: Implications for conservation programs

Despite recent work, uncertainty remains concerning how abiotic and biotic factors affect duckling survival. Additionally, upland habitat characteristics may affect duckling survival rates but this potential relationship has largely been ignored. We evaluated several unresolved hypotheses about causes of mallard (Anas platyrhynchos) duckling survival variation, with an emphasis on assessing effects of managed and remnant natural upland habitats. During 1993–2000, 617 radio-marked females provided information about brood habitat use and duckling survival on 27 sites in prairie Canada. We contrasted a priori and exploratory models that incorporated effects of upland, wetland, weather, female, and brood-related variables on duckling survival rates. Survival was highest for ducklings when a greater proportion of their surrounding landscape (i.e., within a 500-m radius buffer around the brood) was comprised of wetlands characterized by a central expanse of open water and a peripheral ring of flooded emergent vegetation. Cold and wet weather in the first week of life resulted in lower duckling survival. In a post hoc analysis, duckling survival (of older ducklings) was negatively related to increasing proportions of managed hayland. © 2011 The Wildlife Society.     

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi‐precocial Herring Gulls

Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.     

Factors affecting duckling survival of Eiders Somateria mollissima in northeast Scotland

Mortality factors affecting Eider ducklings Somateria mollissima and their impact with respect to age were studied on the Ythan estuary, Aberdeenshire, Scotland. Survival from hatching to fledging averaged 10.4% over 13 years; numbers declined steeply during the first three to four weeks and slowly thereafter. Growth of tagged young varied considerably between broods. Mortality was not correlated with numbers of ducklings or adults.
Predation in 1974 averaged 13.6% per day during the first two weeks, and accounted for most mortality. Herring Gulls Larus argentatus were the principal predators. Predation was 2.5 times as high in rainy, windy weather as in fine, calm weather. Weather, primarily mean rainfall and wind speed, during early life accounted for over 90% of all variation in survival during eight of nine years. Predation was considered the proximate factor in most weather-related mortality. High survival in 1971 was not explained by weather. An estimated 25–40% of mortality during the second week of life was from renal coccidiosis Eimeria somateriae. Reproductive output of the Ythan population probably limits the rate of population growth, but numbers of breeding birds seem to be regulated by other factors acting after fledging.     

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.     

Brood size and body condition in the House Sparrow Passer domesticus: the influence of brooding behaviour

In many bird species, females undergo a marked decline in body condition during the first days of the nestling period. This decline may be because brooding young chicks reduces the time available for foraging. Alternatively, it might be viewed as an adaptive way to reduce flight costs when the food demand of the brood is highest. To test these hypotheses we modified the brooding commitment of House Sparrows Passer domesticus by manipulating brood size to see if changes in time spent brooding affects adult body condition. During the nestling period, females provided on average three times as much brooding as males. Reduced broods received 14% more brooding than large broods and time spent brooding declined with brood size and chick age according to an exponential decay function. Male body condition was unaffected by brood size and remained stable throughout the reproductive period. Body condition of females with enlarged broods decreased gradually during the nestling period, whereas that of females tending reduced broods dropped abruptly and significantly upon hatching. This resulted in females with reduced broods having lower body condition during the first half of the nestling period than those with enlarged broods. The sharp drop in body condition of females with reduced broods coincided with the period that brooding was most intensive. Indeed, female body condition at the end of the nestling period was negatively correlated with the proportion of time they spent brooding during the first half of the nestling period. Thus, the probable lower homeothermic capacities of reduced broods implies a higher brooding commitment for female House Sparrows that, in turn, may reduce their opportunity to forage and consequently also their body condition.     

Cooperation, conflict, and crèching behavior in goldeneye ducks

Crèching behavior, or brood amalgamation, results in offspring being reared by adults other than their genetic parents. Although a variety of hypotheses have been proposed to explain this behavior, most assume either that brood amalgamation is accidental (i.e., nonselected) or that adoption of young is selected for because of social benefits to the young and/or adopting parents. We propose, instead, that brood amalgamation is a function of two separate processes: brood desertion and brood adoption. To examine brood desertion, we develop a graphic model to predict when parents should abandon their young and we test this model experimentally for the Barrow's goldeneye (Bucephala islandica). As predicted, females deserted their offspring when the size of the brood was experimentally reduced. Brood adoption occurred when deserted ducklings joined other broods. However, the success of ducklings in doing so was strongly dependent on the availability of potential host broods and on the age of the recipient broods. Foreign ducklings were readily accepted into young broods (<10 d old) but invariably were rejected from old broods. We could detect no benefits or costs of brood adoption to the host females, contrary to the expectations of a social benefit hypothesis. Our experiments indicate that Crèching behavior is driven by selection on adults to abandon their brood when the benefits of continued investment are outweighed by the reduction in future reproduction and selection on deserted ducklings to join other broods to obtain parental care. Rather than a form of cooperative brood care, Crèching in goldeneyes is perhaps best considered as a form of reproductive parasitism, entailing parent-offspring conflict over brood desertion and intergenerational conflict over adoption of abandoned young.     

Seasonal clutch size decline and individual variation in the timing of breeding are related to female body condition in a non-migratory species, the Upland Goose Chloephaga picta leucoptera

Several studies have shown that waterfowl females in good condition lay larger clutches and start egg-laying earlier in a breeding season. However, most of these studies lack corresponding data on individual condition and timing of breeding in different years. We analysed data on clutch size, egg size, hatching date and female body condition of a non-migratory species, the Upland Goose Chloephaga picta leucoptera, recorded in three seasons on New Island, Falkand Islands. We found a strong seasonal decline in both egg and clutch size. Egg size increased with clutch size. The mean egg volume and total clutch volume increased with female body condition and hatching date was earlier for females in higher body condition. Chicks hatched from nests with higher mean egg volume had higher early body condition. We also compared individual females between different study years and found that individual body condition showed a significant repeatability between years, whereas hatching date did not. While female aging did not systematically affect hatching date, females produced clutches that hatched earlier in years of higher body condition. We could thus show for the first time that the frequently reported negative relationship between female condition and hatching date also holds true on an individual basis. To our knowledge, this is the first study on seasonal clutch size decline in sheldgeese and one of a few in waterfowl comparing condition and timing of breeding within individuals between different years.     

Impact of weather and climate variation on Hoopoe reproductive ecology and population growth

Preserving peripheral populations is a key conservation issue because of the adaptive potential to environmental change they provide for the species as a whole. Yet, peripheral populations are often small and isolated, i.e. more vulnerable to stochastic events and prone to extinction. We studied a peripheral population of Hoopoe (Upupa epops), a rare insectivorous farmland bird, in the Swiss Alps. We first investigated the effect of weather variation on food provisioning to chicks by Hoopoe parents. Second, while accounting for density-dependence, we tested the extent to which breeding success is governed by weather circumstances and assessed the possible consequences of climate variation on population growth. Provisioning rate and provisioned prey biomass were negatively affected by adverse weather (cool, rainy days), were higher in males and also increased with brood size. Much smaller proportions of molecrickets (Gryllotalpa gryllotalpa; the most profitable prey locally, constituting 93% of chicks’ food biomass) were provisioned on days with adverse weather, irrespective of brood size. Rainfall prior to hatching and during the first days of chick life had a negative impact on their survival, and there was a positive effect of temperature on chick survival just before fledging. Reproductive output was negatively affected by precipitation during the hatching period, but was enhanced by warm temperature just before hatching and in the last days before fledging. Our model showed that the variable reproductive output has a strong impact on the population growth: a succession of adverse, rainy springs would cause a rapid decline of the population. This case study confirms that conservation efforts may be obliterated if risks linked to increasing climate variability are not properly accounted for in the management of small peripheral populations.     

Seasonal variation in pre‐fledging survival of lesser scaup Aythya affinis: hatch date effects depend on maternal body mass

Among temperate‐breeding birds, offspring survival and reproductive success are often inversely related to timing of breeding. The mechanisms that produce seasonal declines in offspring survival are not fully understood but may be related to temporal changes in parental quality, environmental quality, or both. We analyzed data for lesser scaup Aythya affinis to evaluate hypothesized effects of parental quality and date on pre‐fledging survival. Maternal quality, as indexed by body mass, did not have an independent effect on offspring survival in this species. Maternal body mass did not decline seasonally and did not have an independent effect on duckling survival. Although we did not detect an independent effect of hatch date on duckling survival, duckling survival declined seasonally for broods raised by lightweight females, indicating an interactive effect of maternal mass and date. We hypothesize that this interaction may be driven by seasonally declining food resources coupled with the influence of female condition on the ability to monopolize food resources or remain attentive to the brood. We also tested morphological predictions of the date hypothesis by examining physical characteristics of ducklings. When corrected for age and size, late‐hatched ducklings tended to have marginally larger digestive systems and smaller leg muscles than did early‐hatched birds. Abundances of intestinal parasites acquired through diet decreased marginally in late‐hatched ducklings. Results for digestive system and parasite infection patterns suggested that later‐hatched broods may shift diets, consistent with a contribution of environmental factors to seasonal variation in offspring survival. Taken together, our results suggest that both female attributes and environmental conditions may influence seasonal patterns of offspring survival in this species.     

Experimental dissociation of individual quality, food and timing of breeding effects on double-brooding in a migratory songbird

Annual reproductive success in many species is influenced by the number of breeding attempts within a season. Although previous studies have shown isolated effects of female quality, food, and timing of breeding on the probability of female birds producing second broods, to our knowledge, none have tested the relative importance of multiple factors and their interactions using simultaneous manipulations within populations of free-living birds. In this study, we show that individual quality and timing of breeding interact to affect the probability of double-brooding in female mountain bluebirds (Sialia currucoides). High-quality females (those that naturally initiated clutches early in the season) were more likely to double-brood, regardless of whether their hatching date was advanced or delayed, whereas later breeding, lower quality females were much less likely to double-brood when their first attempt was delayed. This indicates that annual fecundity of poorer quality (or younger) female bluebirds may be more sensitive to seasonal variation in environmental conditions. In addition, birds that were provided with supplemental food throughout first breeding attempts were more likely to double-brood in one of the study years, suggesting that female bluebirds may be energetically limited in their capacity to initiate a second brood. Females that had their first brood delayed also had a shorter inter-brood interval and were moulting fewer feathers during second broods compared to controls, while females in better condition showed more advanced moult in second breeding attempts. Taken together, our results demonstrate the combined effects of age- or individual quality-mediated energetic trade-offs between current and future reproduction, and between investments in offspring and self-maintenance, on annual fecundity of female birds.     

Factors affecting the probability of double brooding by Southern House Wrens

Seasonal fecundity of birds is influenced by clutch sizes and the number of successful breeding attempts during a breeding season. As such, understanding the factors that determine the decision to initiate multiple broods within a season and the consequences of this reproductive tactic is important. We examined the frequency of double brooding by Southern House Wrens (Troglodytes aedon musculus) in eastern Argentina. We analyzed inter‐ and intraseasonal variation in double brooding and evaluated the effect of weather conditions and laying date on the frequency and occurrence of this behavior. Finally, we assessed the effect of double brooding on the seasonal and lifetime productivity of female Southern House Wrens. During our 8‐year study, we found that ～43% (range = 17–83% each year) of breeding pairs attempted a second brood after successfully raising a first brood. The probability of females having a second brood was affected by the laying date of the first nesting attempt, but was independent of the number of young fledged. About 65% of females that started laying eggs before the first quarter of each breeding season produced a second brood, and this percentage decreased to ～40% after this period. In addition, variation in double‐brooding frequency among years was related to weather conditions, with the proportion of pairs double brooding increasing with increased precipitation early in the breeding season. More precipitation likely contributed to an increase in insect abundance. Although double brooding increased the seasonal and lifetime productivity of female Southern House Wrens, additional study of the survival and fate of fledglings from first and second broods is needed to assess the importance of multi‐brooding in the reproductive success of these wrens.     

Spatial structure and parental aggression in eider broods

The spatial position of young animals within a brood affects their survival, so that marginal individuals are at greater risk of predation. Spatial brood structuring may be caused by differences in offspring size, age, hunger, or active parental manipulation through aggression. Nepotistic manipulation of brood structure would confer fitness benefits for parents accepting nondescendant young. However, insufficient kin recognition has often been considered to preclude such nepotism in birds, particularly in precocial waterfowl. We explored the spatial structure of ducklings within broods of eiders, Somateria mollissima, a seaduck with frequent brood amalgamation. We compared the distribution of ducklings of different origin relative to reference females whose kinship to the ducklings was known. We also observed female aggression towards ducklings, to evaluate the role of parental manipulation of brood structure. We found a nonrandom distribution of ducklings within broods; a female's own young were on average closer to her than unrelated young were. We also found evidence for parental nepotism: whether the brood contained unrelated young was the strongest predictor of female aggression towards ducklings. The spatial position, hatch weight and relative size of ducklings showed no significant correlations with each other, suggesting that active parental manipulation may be needed to explain the observed spatial structure. Our study conflicts with previous anecdotal evidence suggesting that brood amalgamation in eiders results in the disintegration of parent-offspring bonds, preventing parental exploitation of nondescendant young. It also opens up the possibility that the spatial position of ducklings depends on their mother's status in the female dominance hierarchy. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Factors affecting recruitment age and recruitment probability in the Western Gull Larus occidentalis

We studied recruitment age (i.e. the age that a gull bred for the first time) and the probability that an adult reproduced during its life in three cohorts of Western Gulls Larus occidentalis breeding on southeast Farallon Island. We examined the effects of sex, hatching date, hatching order, brood size and annual variation in food supply during the recruitment period. Modal recruitment ages of males and females were 4 and 5 years, respectively, a difference that was significant. None of the factors examined, including hatching date, hatching order and brood size, had a significant effect on recruitment age. The probability of breeding in males who experienced low food supply in their fourth and fifth years was lower than that of males who experienced favourable food supply in their fourth and fifth years. Recruitment probability (the proportion of the sample group that recruited among those available to recruit during a given year) of 4- and 5-year-old males was lower in the food-poor 1983 El Nino than in years of favourable food supply. The only variable affecting recruitment probability of females was age: recruitment probability was greater among 5-year-olds than 4-year-olds. Recruitment age in males was more sensitive to annual variation in food supply, probably because females have more difficulty recruiting due to an excess of adult females in the population and because of the different roles in reproduction in which males are primarily responsible for provisioning chicks and maintaining the territory.     

Maternal developmental stress reduces reproductive success of female offspring in zebra finches

Environmental factors play a key role in the expression of phenotypic traits and life-history decisions, specifically when they act during early development. In birds, brood size is a main environmental factor affecting development. Experimental manipulation of brood sizes can result in reduced offspring condition, indicating that developmental deficits in enlarged broods have consequences within the affected generation. Yet, it is unclear whether stress during early development can have fitness consequences projecting into the next generation. To study such trans-generational fitness effects, we bred female zebra finches, Taeniopygia guttata, whose mothers had been raised in different experimental brood sizes. We found that adult females were increasingly smaller with increasing experimental brood size in which their mother had been raised. Furthermore, reproductive success at hatching and fledging covaried negatively with the experimental brood size in which their mothers were raised. These results illustrate that early developmental stress can have long-lasting effects affecting reproductive success of future generations. Such trans-generational effects can be life-history responses adapted to environmental conditions experienced early in life.     

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993–1996). The sex of 465 nestlings from 169 broods was determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in any year and the variance in brood sex ratios did not deviate from the binomial distribution. Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size. The sex ratio of broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience. However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding of fairy martin life history and breeding ecology.     

The influence of environmental conditions on immune responses, morphology and recapture probability of nestling house martins (Delichon urbica)

Nestling birds produced later in the season are hypothesized to be of poor quality with a low probability of survival and recruitment. In a Spanish population of house martins (Delichon urbica), we first compared reproductive success, immune responses and morphological traits between the first and the second broods. Second, we investigated the effects of an ectoparasite treatment and breeding date on the recapture rate the following year. Due probably to a reverse situation in weather conditions during the experiment, with more rain during rearing of the first brood, nestlings reared during the second brood were in better condition and had stronger immune responses compared with nestlings from the first brood. Contrary to other findings on house martins, we found a similar recapture rate for chicks reared during the first and the second brood. Furthermore, ectoparasitic house martin bugs had no significant effect on the recapture rate. Recaptured birds had similar morphology but higher immunoglobulin levels when nestlings compared with non-recaptured birds. This result implies that a measure of immune function is a better predictor of survival than body condition per se.     

Post-hatching parental care behaviour and hormonal status in a precocial bird

In birds, the link between parental care behaviour and prolactin release during incubation persists after hatching in altricial birds, but has never been precisely studied during the whole rearing period in precocial species, such as ducks. The present study aims to understand how changes in parental care after hatching are related to circulating prolactin levels in mallard hens rearing ducklings. Blood was sampled in hens over at least 13 post-hatching weeks and the behaviour of the hens and the ducklings was recorded daily until fledging. Contacts between hens and the ducklings, leadership of the ducklings and gathering of them steadily decreased over post-hatching time. Conversely, resting, preening and agonistic behaviour of hens towards ducklings increased. Plasma prolactin concentrations remained at high levels after hatching and then fell after week 6 when body mass and structural size of the young were close to those of the hen. Parental care behaviour declined linearly with brood age, showed a disruption of the hen-brood bond at week 6 post-hatching and was related to prolactin concentration according to a sigmoid function. Our results suggest that a definite threshold in circulating prolactin is necessary to promote and/or to maintain post-hatching parental care in ducks.