Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

Breeding performance and survival in the peregrine falcon Falco peregrinus support an age‐related competence improvement hypothesis mediated via an age threshold

We studied the effects of age on breeding performance and survival probability in a peregrine falcon population, using data from a long term monitoring programme (carried out over 16 yr), in which we were able to identify individual birds. We compared the breeding performance and survival of yearling breeders, first‐time adult breeders and adult breeders. We found significant differences in breeding performance but not in survival. Yearling breeders had lower breeding success than older individuals but the breeding performance of inexperienced adults did not differ from that of experienced adults. We did not find changes in terminal breeding success since peregrines in their last year of life sustained the performance levels shown in previous years although with increasing variability. We found no evidence that attempting to breed affected survival probability in any age group. We argue that differences in breeding performance are related to age, not to breeding experience, and that there is an age threshold, coincident with the development of adult plumage, after which breeding performance is not affected either by age or experience. Peregrines that start breeding as yearlings are likely to have greater lifetime reproductive success than birds entering the breeding pool as adults. Consequently, such birds may represent a set of high quality individuals. Our results support the age‐related competence improvement hypothesis as being the relevant explanation for the increase in breeding performance with age.     

Age-specific survival and reproductive performances in fur seals: evidence of senescence and individual quality

Life history theory hypothesises that breeding events induce reproductive costs that may vary among individuals. However, the growing number of studies addressing this question are taxonomically biased, therefore impeding the generalisation of this hypothesis, especially with regard to marine top predators. This study investigated age‐related survival and breeding performances in subantarctic fur seal (Arctocephalus tropicalis) females from Amsterdam Island, southern Indian Ocean. Using multistate capture–recapture models on data obtained from known‐age tagged females over eight consecutive years, we tested for evidence of senescence, individual quality, and reproductive costs in terms of future survival and fecundity. Adult female yearly survival appeared high and constant throughout time. While a two age‐class model was preferred in non‐breeders, breeding females exhibited three age classes with a maximum survival for the prime‐age class (7–12 years). Survival and reproductive probabilities decreased from 13 years onward, suggesting senescence in this population. Survival was lower for non‐breeders than for breeders, among both prime‐aged (0.938 vs 0.982) and older (0.676 vs 0.855) females. Furthermore, non‐breeders exhibited higher probabilities of being non‐breeders the following year than did breeders (0.555 vs 0.414). Such results suggest consistency in female breeding performance over years, supporting the hypothesis that non‐breeding tend to occur among lower quality individuals rather than representing an alternative strategy to enhance residual reproductive value. However, the high proportion of females that did not breed during two consecutive years, and the lower probability of being a successful breeder after a greater reproductive effort confirmed the existence of reproductive costs, especially during the second half of the lactation. These results also suggest that younger age‐classes included a higher proportion of lower quality individuals, which are likely to face higher costs of reproduction. Such hypotheses lead to consider the first breeding event as a filter generating a within‐cohort selection process in females.     

Population dynamics in a long-lived seabird: I. Impact of breeding activity on survival and breeding probability in unbanded king penguins

Understanding the trade-off between current reproductive effort, future survival and future breeding attempts is crucial for demographic analyses and life history studies. We investigated this trade-off in a population of king penguins (Aptenodytes patagonicus) marked individually with transponders using multistate capture-recapture models. This colonial seabird species has a low annual proportion of non-breeders (13%), despite a breeding cycle which lasts over 1 year. To draw inferences about the consequences of non-breeding, we tested for an effect of reproductive activity on survival and on the probability of subsequent breeding. We found that birds non-breeding in year t show the same survival rate as breeders (two-states analysis: breeding and non-breeding). However, breeders had a lower probability of breeding again the following year. This negative phenotypic correlation suggests the existence of reproductive costs affecting future breeding probability, but it might also be strengthened by late arrival for courtship in year t. A three-state analysis including breeding success revealed that failed breeders in year t have a lower probability to reproduce successfully in year t + 1 than non-breeders in year t, providing some evidence for the existence of reproductive costs. Moreover, successful breeders showed higher survival probability. This positive phenotypic correlation between current reproduction and subsequent survival supports the hypothesis of an heterogeneity in individual quality. Males breeding in year t had a lower probability to breed again in year t + 1 than females, suggesting higher reproductive costs for this sex. Such additional costs might be due to higher male parental investment in the final phase of chick-rearing, which also delays the arrival of males in year t + 1, and decreases their breeding probability. Our study is the first to explore the breeding biology and the demography of penguins without the disturbance of flipper-bands.     

Best squirrels trade a long life for an early reproduction

Age at primiparity plays a crucial role in population dynamics and life-history evolution. Long-term data on female North American red squirrels were analysed to study the fitness consequences of delaying first reproduction. Early breeders were born earlier, had a higher breeding success and achieved a higher lifetime reproductive success than females who delayed their first reproduction, which suggests a higher quality of early breeders. However, early breeders had similar mass when tagged, and similar number of food caches available at one year of age as late breeders. Nevertheless, we found evidence of survival costs of early primiparity. Early breeders had a lower survival between one and two years of age than late breeders and a lower lifespan. Our study points out that two reproductive tactics co-occurred in this population: a tactic based on early maturity at the cost of a lower survival versus a tactic based on delayed maturity and long lifespan. High quality individuals express the most profitable tactic by breeding early whereas low quality individuals do the best of a bad job by delaying their first reproduction.     

Experience versus effort: what explains dynamic heterogeneity with respect to age?

Age‐related patterns of survival and reproduction have been explained by accumulated experience (‘experience hypothesis’), increased effort (‘effort hypothesis’), and intrinsic differences in phenotypes (‘selection hypothesis’). We examined the experience and effort hypotheses using a 40‐year data set in a population of Leach's storm‐petrels Oceanodroma leucorhoa, long‐lived seabirds for which the effect of phenotypic variation has been previously demonstrated. Age was quantified by time since recruitment (‘breeding age’). The best model of adult survival included a positive effect of breeding age (1, 2, 3+ years), sex (male > female), and year. Among‐individuals variation (fixed heterogeneity) accounted for 31.6% of the variance in annual reproductive success. We further examined within‐individual patterns in reproductive success (dynamic heterogeneity) in the subset of individuals with at least five breeding attempts. Three distinct phases characterized reproductive success – early increase, long asymptotic peak, late decline. No effect of early reproductive output on longevity was found, however, early success was positively correlated with lifetime reproductive success. Reproductive success was lower earlier than later in life. Among the few natally philopatric individuals in the population, age of first breeding had no effect on longevity, lifetime reproductive success, or early reproductive success. No support for the effort hypothesis was found in this population. Instead, age‐specific patterns of survival and reproduction in these birds are best explained by the experience hypothesis over and above the effect of intrinsic differences among individuals.     

Environmental variation and experience-related differences in the demography of the long-lived black-browed albatross

1. It has been largely demonstrated that demographic performances of animals increase with age or experience as a result of an improvement of foraging skills, an increasing reproductive effort or a selection process. However, little is known about the age or experience-related response of populations to environmental variations. Theoretical studies consider that age-related variations of the performances are greater under more restricting conditions, but this has rarely been tested. 2. We tested this hypothesis on a long-lived species, black browed albatross Thalassarche melanophrys Temminck, using a long-term capture-mark-recapture data set. We investigated the responses of a population to climate, by studying the effects of climatic factors and breeding experience on survival and breeding success. 3. First-time breeders appear to be poorer performers compared with experienced adults, with lower reproductive success and lower survival. In addition, interannual variations of demographic traits were partly explained by climatic indices, reflecting environmental variations. The survival probability of black-browed albatrosses varied with experience and climate, and differences being greater under harsh conditions. By contrast, the reproductive success of inexperienced individuals was affected by climatic fluctuations in the same way as the experienced ones. 4. First breeding event acts as a strong selective process on the highly heterogeneous class of inexperienced individuals, suggesting the increase in survival and breeding success with experience may mainly reflect a reduction in the heterogeneity among individual qualities.     

Improvement of reproductive performance with age and breeding experience depends on recruitment age in a long‐lived seabird

Reproductive success increases with age in many species, however, the underlying mechanisms remain unclear and several hypotheses have been proposed to explain age‐related improvements in reproductive output. In this contribution we investigated the effects of age, recruitment age, breeding experience and sex on reproductive performance during the early breeding career in the common tern Sterna hirundo using long‐term individual‐based data. We used measurements of performance, which spanned the entire breeding process: clutch size, hatching success, fledging success and fledglings per pair. Longitudinal analyses within individuals showed a clear increase with age in all performance measures. Furthermore, a significant change in reproductive performance was found between first time‐ and experienced breeders. Recruitment age had a strong influence on hatching and fledging success: two‐year‐old recruits had significantly lower reproductive success than birds which recruited at older ages, but the increase in breeding performance with experience was stronger in young recruits. Comparing age and experience effects, age effects were more pronounced during the first breeding attempts, whereas experience effects were also visible in subsequent breeding attempts. The degree of intra‐individual improvements in reproductive performance is due to a complex interplay of age at first breeding and experience. The results strongly support the constraint hypothesis.     

Stratification based on reproductive state reveals contrasting patterns of age-related variation in demographic parameters in the kittiwake

Heterogeneity in individual quality can be a major obstacle when interpreting age‐specific variation in life‐history traits. Heterogeneity is likely to lead to within‐generation selection, and patterns observed at the population level may result from the combination of hidden patterns specific to subpopulations. Population‐level patterns are not relevant to hypotheses concerning the evolution of age‐specific reproductive strategies if they differ from patterns at the individual level. We addressed the influence of age and a variable used as a surrogate of quality (yearly reproductive state) on survival and breeding probability in the kittiwake. We found evidence of an effect of age and quality on both demographic parameters. Patterns observed in breeders are consistent with the selection hypothesis, which predicts age‐related increases in survival and traits positively correlated with survival. Our results also reveal unexpected age effects specific to subgroups: the influence of age on survival and future breeding probability is not the same in nonbreeders and breeders. These patterns are observed in higher‐quality breeding habitats, where the influence of extrinsic factors on breeding state is the weakest. Moreover, there is slight evidence of an influence of sex on breeding probability (not on survival), but the same overall pattern is observed in both sexes. Our results support the hypothesis that age‐related variation in demographic parameters observed at the population level is partly shaped by heterogeneity among individuals. They also suggest processes specific to subpopulations. Recent theoretical developments lay emphasis on integration of sources of heterogeneity in optimization models to account for apparently “sub‐optimal” empirical patterns. Incorporation of sources of heterogeneity is also the key to investigation of age‐related reproductive strategies in heterogeneous populations. Thwarting “heterogeneity's ruses” has become a major challenge: for detecting and understanding natural processes, and a constructive confrontation between empirical and theoretical studies.     

Age-related change in breeding performance in early life is associated with an increase in competence in the migratory barn swallow Hirundo rustica

1. We investigated age-related changes in two reproductive traits (laying date and annual fecundity) in barn swallows Hirundo rustica L. using a mixed model approach to di-stinguish among between- and within-individual changes in breeding performance with age. 2. We tested predictions of age-related improvements of competence (i.e. constraint hypothesis) and age-related progressive disappearance of poor-quality breeders (i.e. selection hypothesis) to explain age-related increase in breeding performance in early life. 3. Reproductive success increased in early life, reaching a plateau at middle age (e.g. at 3 years of age) and decreasing at older age (> 4 years). Age-related changes in breeding success were due mainly to an effect of female age. 4. Age of both female and male affected timing of reproduction. Final linear mixed effect models (LME) for laying date included main and quadratic terms for female and male age, suggesting a deterioration in reproductive performance at older age for both males and females. 5. We found evidence supporting the constraints hypothesis that increases in competence within individuals, with ageing being the most probable cause of the observed increase in breeding performance with age in early life. Two mechanisms were implicated: (1) advance in male arrival date with age provided middle-aged males with better access to mates. Yearling males arrived later to the breeding grounds and therefore had limited access to high-quality mates. (2) Breeding pairs maintaining bonds for 2 consecutive years (experienced pairs) had higher fecundity than newly formed inexperienced breeding pairs. 6. There was no support for the selection hypothesis because breeding performance was not correlated with life span. 7. We found a within-individual deterioration in breeding and migratory performance (arrival date) in the oldest age-classes consistent with senescence in these reproductive and migratory traits.     

Age-related improvement in reproductive performance in a long-lived raptor: a cross-sectional and longitudinal study

In numerous iteroparous species, mean fecundity increases with age. Such improvement has been explained by: a) progressive removal of inferior breeders from the breeding population (selection‐hypothesis); b) delayed breeding of higher‐quality phenotypes (delayed‐breeder‐hypothesis); c) longitudinal enhancement of skills associated with age per se (age‐hypothesis); d) progressive improvement in the capability to conduct specific tasks facilitated by accumulated experience (breeding‐experience‐hypothesis); and e) increasing parental investment promoted by declining residual reproductive values (restraint‐hypothesis). To date, there have been few comprehensive tests of these hypotheses. Here, we provide such a study using a long‐term dataset on a long‐lived raptor, the black kite Milvus migrans (maximum lifespan 23 yr). Kites delayed breeding for 1–7 yr and all measures of breeding performance increased linearly or quadratically up to 11 yr of age. There was no support for the delayed‐breeder‐hypothesis: superior phenotypes did not delay breeding longer. Superior breeders were retained longer in the breeding population, consistent with the selection‐hypothesis. All measures of breeding performance increased longitudinally within individuals, supporting the age‐hypothesis, while some of them increased with accumulated previous experience, supporting the breeding‐experience‐hypothesis. Some analyses suggested the existence of trade‐offs between reproduction in the early years of life and subsequent survival, partially supporting the restraint‐hypothesis. The pattern of age‐related improvements in breeding rates observed at the population‐level could be ascribed to the combined effect of the progressive removal of inferior phenotypes from the breeding population and the individual‐level lack of specific skills which are progressively acquired with time and experience. It was also compatible with a longitudinal increase in reproductive investment. Results from previous studies suggest that different mechanisms may operate in different species and that age‐related improvements in reproduction may be frequently promoted by the complex interplay between longitudinal improvements and changes in the relative frequency of productive phenotypes in the breeding population.     

Age-specific reproductive success: evidence for the selection hypothesis

Age-specific reproductive success has been demonstrated in many species. Three hypotheses have been raised to explain this general phenomenon: the experience hypothesis based on age-specific reproductive experience, the effort hypothesis based on age-specific reproductive effort, and the selection hypothesis based on progressive disappearance of phenotypes due to variation in individual productivity and survival. We used data from a long-term study of Leach's storm-petrels (Oceanodroma leucorhoa) to present a single test of mutually exclusive predictions about the relationship between early breeding success and longevity. There should be no correlation between early breeding success and longevity under the experience hypothesis, a negative correlation under the effort hypothesis, and a positive correlation under the selection hypothesis. We found a significant (P < 0.0001) positive relationship between success in the first two breeding attempts and longevity in this population of long-lived seabirds, strongly suggesting that low-productivity parents were also less likely to survive early breeding. These data provide some of the strongest support to date for the selection hypothesis.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.     

Reproductive compensation

The reproductive compensation hypothesis says that individuals constrained by ecological or social forces to reproduce with partners they do not prefer compensate for likely offspring viability deficits. The reproductive compensation hypothesis assumes that (i) pathogens and parasites evolve more rapidly than their hosts, (ii) mate preferences predict variation in health and viability of offspring, (iii) social and ecological factors keep some individuals from mating with their preferred partners (some are constrained to mate with partners they do not prefer), (iv) all individuals may be induced to compensate, so that (v) variation in compensation is due to environmental and developmental factors affecting between-individual abilities to express compensatory mechanisms. Selection favouring compensation may act through variation in prezygotic physiological mechanisms, zygotic mechanisms, or parental care to eggs or young that enhance offspring health, increasing the likelihood that some offspring survive to reproductive age, often at a survival cost to the parents. Compensation may be through increased number of eggs laid or offspring born, a compensatory effort working during a single reproductive bout that sometimes will match the number of offspring surviving to reproductive age produced by unconstrained parents during the same bout. The reproductive compensation hypothesis therefore predicts trade-offs in components of fitness for breeders, such that parents constrained to mating with a nonpreferred partner, but who compensate sometimes match their current productivity (number of offspring at reproductive age) to unconstrained parents (those breeding with their preferred partners), and, when all else is equal, die faster than unconstrained parents. The reproductive compensation hypothesis emphasizes that reproductive competition is not just between constrained and unconstrained individuals, but also among constrained individuals who do and do not compensate. The reproductive compensation hypothesis may thus explain previously unexplained between-population and within-population, between-individual variation in reproductive success, survival, physiology and behaviour.     

Attendance off breeders and prospectors reflects the quality off colonies in the Kittiwake Rissa tridactyla

Attendance of breeders and prospectors (prebreeders, failed breeders and non-breeding adults) was studied from the time of hatching to the end of the breeding season in parts of two neighbouring Kittiwake Rissa tridactyla colonies in Brittany (France). Many clutches in one subcolony had been predated by a Carrion Crow Corvus corone , leading to a lower breeding success. Attendance of breeders and prospecting activity were greater in the more successful colony, and the number of breeding pairs increased in the following year in this colony only. Local reproductive success and social behaviour contributed to the attraction exerted by a colony. The demographic trend was explained by a performance-based conspecific attraction mechanism, acting through the effect of local reproductive success on recruitment, non-breeding and fidelity. Study of breeding success and prospecting activity allowed both evaluation of the quality of a colony and prediction of local demographic trends for the following year.     

Breeding dispersal strategies following reproductive failure explain low apparent survival of immigrant Temminck's stints

In some animal populations, immigrants have lower survival than philopatric individuals. Costs of dispersal or low phenotypic quality of dispersers may explain the pattern. However, apparent adult survival estimates, which describe real survival combined with site fidelity cannot be separated from permanent emigration. Thus, heterogeneity in breeding dispersal propensities of immigrants and philopatrics can bias fitness correlates of dispersal. Differences in breeding dispersal propensities may be caused by different strategies in response to environmental cues inducing dispersal, such as reproductive success. In such cases, the reported differences between immigrants and philopatric individuals may not reflect true variation in survival. We studied whether dispersal status specific apparent adult survival is associated with reproductive success in a Temminck's stint Calidris temminckii population. We analysed two long term capture–recapture datasets characterised by low and high nest predation levels. Philopatric individuals had higher apparent adult survival than immigrants in both datasets and the difference was highlighted during the high nest predation period. By contrasting return rates between successful and unsuccessful breeders as a proxy for dispersal, we found that unsuccessful immigrants breeding for the first time dispersed more likely than successful immigrants, but such a pattern was not found among philopatric individuals. Our results support the hypothesis that immigrant and philopatric individuals have different breeding dispersal strategies following reproductive failure and that their apparent adult survival differences are at least partly explained by different breeding dispersal propensities. Our results also suggest that the recent decline of the study population reflects a multiple response to increased nest predation through decreased local recruitment and increased emigration.     

Long- and short-term influence of environment on recruitment in a species with highly delayed maturity

Short-term effects of environmental perturbations on various life history traits are reasonably well documented in birds and mammals. But, in the present context of global climate change, there is a need to consider potential long-term effects of natal conditions to better understand and predict the consequences of these changes on population dynamics. The environmental conditions affecting offspring during their early development may determine their lifetime reproductive performance, and therefore the number of recruits produced by a cohort. In this study, we attempted to link recruitment to natal and recent (previous year) conditions in the long-lived black-browed albatross (Thalassarche melanophrys) at Kerguelen Islands. The environmental variability was described using both climatic variables over breeding (sea surface temperature anomaly) and non-breeding grounds (Southern Oscillation index), and variables related to the colony (breeding success and colony size). Immature survival was linked to the breeding success of the colony in the year of birth, which was expected to reflect the average seasonal parental investment. At the cohort level, this initial mortality event may act as a selective filter shaping the number, and presumably the quality (breeding frequency, breeding success probability), of the individuals that recruit into the breeding population. The decision to start breeding was strongly structured by the age of the individuals and adjusted according to recent conditions. An effect of natal conditions was not detected on this parameter, supporting the selection hypothesis. Recruitment, as a whole, was thus influenced by a combination of long- and short-term environmental impacts. Our results highlight the complexity of the influence of environmental factors on such long-lived species, due to the time-lag (associated with a delayed maturity) between the impact of natal conditions on individuals and their repercussion on the breeding population.     

Age-related fledgling production in great cormorants Phalacrocorax carbo: influence of individual competence and disappearance of phenotypes

Age-related reproductive performance of great cormorants Phalacrocorax carbo sinensis was studied in a tree nesting colony in Denmark in relation to age-related improvements of competence and progressive disappearance of phenotypes. Within-individual changes in fledgling production were measured, and cross-sectional analyses were applied. The within-individual analyses showed that competence improved with age and/or that individuals showed restraint to optimize their reproductive effort. The within-individual improvements were three to six times higher among individuals that survived and returned to breed beyond the fourth breeding attempt than among individuals disappearing from the breeding population before the fourth breeding attempt. Taking this into account the within-individual improvements explained 70–90% of the age-effect observed in the population over the youngest ages. Effects of breeding experience were significant for females, but only within the group of individuals that were present in the breeding population beyond the age of five years. In males, improvements arose because of unknown factors related to age. Individual great cormorants that bred beyond the age of five years had higher reproductive success, on average, than birds disappearing from the breeding population earlier in life. This supports the differential survival hypothesis. However, the effect on the population mean was partly counterbalanced by late recruitment of other inferior breeders. It is concluded that the enhancement in fledgling production with increasing age was primarily an effect of age-related improvements of competence and secondly an effect of progressive disappearance of phenotypes.     

Evaluation of reproductive costs for Weddell seals in Erebus Bay, Antarctica

1. Organisms balance current reproduction against future survival and reproduction, which results in life-history trade-offs. These trade-offs are also known as reproductive costs and may represent significant factors shaping life-history strategy for many species. 2. Using multistate mark-resight models and 26 years of mark-resight data (1979-2004), we estimated the costs of reproduction to survival and reproductive probabilities for Weddell seals in Erebus Bay, Antarctica and evaluated whether this species either conformed to the 'prudent parent' reproductive strategy predicted by life-history theory for long-lived mammals or alternatively, incurred costs to survival in order to reproduce in a variable environment (flexible-strategy hypothesis). 3. Results strongly supported the presence of reproductive costs to survival (mean annual survival probability was 0.91 for breeders vs. 0.94 for nonbreeders), a notable difference for a long-lived mammal, demonstrating that investment in reproduction does result in a cost to survival for Weddell seals, contrary to the prudent parent hypothesis. 4. Reproductive costs to subsequent reproductive probabilities were also present for first-time breeders (mean probability of breeding the next year was 31.3% lower for first-time breeders than for experienced breeders), thus supporting our prediction of the influence of breeding experience. 5. We detected substantial annual variation in survival and breeding probabilities. Breeding probabilities were negatively influenced by summer sea-ice extent, whereas weak evidence suggested that survival probabilities were affected more by winter sea-ice extent, and the direction of this effect was negative. However, a model with annual variation unrelated to any of our climate or sea-ice covariates performed best, indicating that further study will be needed to determine the appropriate mechanism or combination of mechanisms underlying this annual variation.     

What shall I do now? State-dependent variations of life-history traits with aging in Wandering Albatrosses

Allocation decisions depend on an organism's condition which can change with age. Two opposite changes in life‐history traits are predicted in the presence of senescence: either an increase in breeding performance in late age associated with terminal investment or a decrease due to either life‐history trade‐offs between current breeding and future survival or decreased efficiency at old age. Age variation in several life‐history traits has been detected in a number of species, and demographic performances of individuals in a given year are influenced by their reproductive state the previous year. Few studies have, however, examined state‐dependent variation in life‐history traits with aging, and they focused mainly on a dichotomy of successful versus failed breeding and non‐breeding birds. Using a 50‐year dataset on the long‐lived quasi‐biennial breeding wandering albatross, we investigated variations in life‐history traits with aging according to a gradient of states corresponding to potential costs of reproduction the previous year (in ascending order): non‐breeding birds staying at sea or present at breeding grounds, breeding birds that failed early, late or were successful. We used multistate models to study survival and decompose reproduction into four components (probabilities of return, breeding, hatching, and fledging), while accounting for imperfect detection. Our results suggest the possible existence of two strategies in the population: strict biennial breeders that exhibited almost no reproductive senescence and quasi‐biennial breeders that showed an increased breeding frequency with a strong and moderate senescence on hatching and fledging probabilities, respectively. The patterns observed on survival were contrary to our predictions, suggesting an influence of individual quality rather than trade‐offs between reproduction and survival at late ages. This work represents a step further into understanding the evolutionary ecology of senescence and its relationship with costs of reproduction at the population level. It paves the way for individual‐based studies that could show the importance of intra‐population heterogeneity in those processes.