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.     

Female red squirrels fit Williams' hypothesis of increasing reproductive effort with increasing age

Williams predicted that reproductive effort should increase as individuals age and their reproductive value declines. This simple prediction has proven difficult to test because conventional measures of energy expenditure on reproduction may not be a true reflection of reproductive effort. We investigated age-specific variation in female reproductive effort in a stable population of North American red squirrels where energy expenditure on reproduction is likely to reflect actual reproductive effort. We used seven measures of reproductive effort spanning conception to offspring weaning. We found that females completed growth by age 3 and that reproductive value decreased after this age likely because of reproductive and survival senescence. We therefore, predicted that reproductive effort would increase from age 3 onwards. The probability of breeding, litter mass at weaning, and likelihood of territory bequeathal were all lower for 1- and 2-year-old females than for females older than 3 years, the age at which growth is completed. That growing females are faced with additional energetic requirements might account for their lower allocation to reproduction as compared with older females. The probability of attempting a second reproduction within the same breeding season and the propensity to bequeath the territory to juveniles increased from 3 years of age onwards, indicating an increase in reproductive effort with age. We think this increase in reproductive effort is an adaptive response of females to declining reproductive values when ageing, thereby supporting Williams' prediction.     

Trade-off between current reproductive effort and delay to next reproduction in the leatherback sea turtle

The trade-off between current and future reproduction plays an important role in demographic analyses. This can be revealed by the relationship between the number of years without reproduction and reproductive investment within a reproductive year. However, estimating both the duration between two successive breeding season and reproductive effort is often limited by variable recapture or resighting effort. Moreover, a supplementary difficulty is raised when nonbreeder individuals are not present sampling breeding grounds, and are therefore unobservable. We used capture–recapture (CR) models to investigate intermittent breeding and reproductive effort to test a putative physiological trade-off in a long-lived species with intermittent breeding, the leatherback sea turtle. We used CR data collected on breeding females on Awa:la-Ya:lima:po beach (French Guiana, South America) from 1995 to 2002. By adding specific constraints in multistate (MS) CR models incorporating several nonobservable states, we modelled the breeding cycle in leatherbacks and then estimated the reproductive effort according to the number of years elapsed since the last nesting season. Using this MS CR framework, the mean survival rate was estimated to 0.91 and the average resighting probability to 0.58 (ranged from 0.30 to 0.99). The breeding cycle was found to be limited to 3 years. These results therefore suggested that animals whose observed breeding intervals are greater than 3 years were most likely animals that escaped detection during their previous nesting season(s). CR data collected in 2001 and 2002 allowed us to compare the individual reproductive effort between females that skipped one breeding season and females that skipped two breeding seasons. These inferences led us to conclude that a trade-off between current and future reproduction exists in leatherbacks nesting in French Guiana, likely linked to the resource provisioning required to invest in 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.     

Reproductive effort and future parental competitive ability: A nest box removal experiment

The life history trade‐off between current and future reproduction is a theoretically well‐established concept. However, empirical evidence for the occurrence of a fitness cost of reproduction is mixed. Evidence indicates that parents only pay a cost of reproduction when local competition is high. In line with this, recent experimental work on a small passerine bird, the Great tit (Parus major) showed that reproductive effort negatively affected the competitive ability of parents, estimated through competition for high quality breeding sites in spring. In the current study, we further investigate the negative causal relationship between reproductive effort and future parental competitive ability, with the aim to quantify the consequences for parental fitness, when breeding sites are scarce. To this end, we (a) manipulated the family size of Great tit parents and (b) induced severe competition for nest boxes among the parents just before the following breeding season by means of a large‐scale nest box removal experiment. Parents increased their feeding effort in response to our family size manipulation and we successfully induced competition among the parents the following spring. Against our expectation, we found no effect of last season's family size on the ability of parents to secure a scarce nest box for breeding. In previous years, if detected, the survival cost of reproduction was always paid after midwinter. In this year, parents did pay a survival cost of reproduction before midwinter and thus before the onset of the experiment in early spring. Winter food availability during our study year was exceptionally low, and thus, competition in early winter may have been extraordinarily high. We hypothesize that differences in parental competitive ability due to their previous reproductive effort might have played a role, but before the onset of our experiment and resulted in the payment of the survival cost of reproduction.     

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.     

Experimental evidence of long‐term reproductive costs in a colonial nesting seabird

Trade‐offs between current and future reproduction are central to the evolution of life histories. Experiments that manipulate brood size provide an effective approach to investigating future costs of current reproduction. Most manipulative studies to date, however, have addressed only the short‐term effects of brood size manipulation. Our goal was to determine whether survival or breeding costs of reproduction in a long‐lived species manifest beyond the subsequent breeding season. To this end, we investigated long‐term survival and breeding effects of a multi‐year reproductive cost experiment conducted on black‐legged kittiwakes Rissa tridactyla, a long‐lived colonial nesting seabird. We used multi‐state capture–recapture modeling to assess hypotheses regarding the role of experimentally reduced breeding effort and other factors, including climate phase and colony size and productivity, on future survival and breeding probabilities during the 16‐yr period following the experiment. We found that forced nest failures had a positive effect on breeding probability over time, but had no effect on long‐term survival. This apparent canalization of survival suggests that adult survival is the most important parameter influencing fitness in this long‐lived species, and that adults should pay reproductive costs in ways that do not compromise this critical life history parameter. When declines in adult survival rate are observed, they may indicate populations of conservation concern.     

Early reproductive investment,senescence and lifetime reproductive success in female Asian elephants

The evolutionary theory of senescence posits that as the probability of extrinsic mortality increases with age, selection should favour early‐life over late‐life reproduction. Studies on natural vertebrate populations show early reproduction may impair later‐life performance, but the consequences for lifetime fitness have rarely been determined, and little is known of whether similar patterns apply to mammals which typically live for several decades. We used a longitudinal dataset on Asian elephants (Elephas maximus) to investigate associations between early‐life reproduction and female age‐specific survival, fecundity and offspring survival to independence, as well as lifetime breeding success (lifetime number of calves produced). Females showed low fecundity following sexual maturity, followed by a rapid increase to a peak at age 19 and a subsequent decline. High early life reproductive output (before the peak of performance) was positively associated with subsequent age‐specific fecundity and offspring survival, but significantly impaired a female's own later‐life survival. Despite the negative effects of early reproduction on late‐life survival, early reproduction is under positive selection through a positive association with lifetime breeding success. Our results suggest a trade‐off between early reproduction and later survival which is maintained by strong selection for high early fecundity, and thus support the prediction from life history theory that high investment in reproductive success in early life is favoured by selection through lifetime fitness despite costs to later‐life survival. That maternal survival in elephants depends on previous reproductive investment also has implications for the success of (semi‐)captive breeding programmes of this endangered species.     

Nest predation reduces benefits to early clutch initiation in northern cardinals Cardinalis cardinalis

Life history theory and empirical studies suggest that early breeding confers higher reproductive success, but the extent to which this advantage can be generalized to human‐dominated systems and across species is less well understood. We studied the fitness consequences of clutch initiation for 181 female northern cardinals Cardinalis cardinalis and 1228 nests in forests within urban and rural landscapes of Ohio, USA between 2004–2007. Cardinals that bred earlier made significantly more nesting attempts, but cumulative number of young fledged was similar to that of later‐breeding individuals. The expected number of fledglings produced per successful nest was unrelated to date and remained ~1.8 fledglings across the season, despite the fact that nest survival rates improved dramatically as the season progressed. Because the probability of resighting breeding individuals in subsequent years was unrelated to first clutch initiation date, we have no evidence that clutch initiation affected adult survival. The absence of a clear benefit to early breeding appears to be a consequence of high rates of nest predation early in the breeding season.     

The adaptive value of energy storage and capital breeding in seasonal environments

Timing of reproduction in a seasonal cycle is a life history trait with important fitness consequences. Capital breeders produce offspring from stored resources and, by decoupling feeding and reproduction, may bend the constraints caused by seasonality in food or predation. Income breeders, on the other hand, produce offspring from concurrent food intake, with the disadvantage of less flexibility, but with high efficiency and no inventory costs of carrying stores. Here, we assess relative profitability of capital and income breeding in herbivorous zooplankton inhabiting seasonal, high-latitude environments. We apply a state-dependent life history model where reproductive values are used to optimise energy allocation and diapause strategies over the year. Three environmental scenarios were modelled: an early, an intermediate, and a late feeding season. We found that capital breeding was most important in the early season. Capital breeding ranged from 7–9% of the eggs produced but, because of the high reproductive value of early eggs, capital breeding ranged from 9–30% when measured in terms of reproductive value. The main benefit of capital breeding was reproduction prior to the feeding season – when the reproductive value of an egg peaked. In addition, capital breeding was also used to increase egg production rates at times of income breeding. For individuals born late in the season the model predicted a two-year cycle instead of the typical annual life cycle. These individuals could then reap the benefits of early reproduction and capital breeding in their second year instead of income breeding late in the first year. We emphasize the importance of evaluating reproductive strategies such as capital and income breeding from a complete life cycle perspective. In particular, knowing the seasonality in offspring fitness is essential to appreciate evolutionary and population-level consequences of capital breeding.     

Life history patterns,individual heterogeneity,and density dependence in breeding common goldeneyes of the northern boreal forest

Life history patterns and their associated tradeoffs influence population dynamics, as they determine how individuals allocate resources among competing demographic traits. Here we examined life history strategies in common goldeneyes Bucephala clangula (hereafter goldeneye), a cavity‐nesting sea duck, in the northern boreal forest of interior Alaska, USA. We used multistate capture–mark–recapture models to estimate adult survival, breeding probability, first‐year survival, and recruitment probability using a long‐term nest box study (1997–2010). We detected annual variation in adult survival, which varied from 0.74 ± 0.12 (SE) to 0.93 ± 0.06. In contrast, breeding probability remained relatively high and invariant (0.84 ± 0.11) and was positively related to individual nest success the year prior. Nonbreeding individuals in one year were more likely to remain a nonbreeder, than attempt to breed the following year. First‐year survival decreased with smaller residual duckling mass and larger brood sizes. Probability of recruitment into the breeding population conditioned on survival was constant during the study (0.96 ± 0.06), and did not vary among ages 2–5 yr‐old. Overall, goldeneyes exhibited high, but somewhat variable, adult survival, and high breeding and recruitment probabilities, which is consistent with observed patterns in bet‐hedging species that breed annually in high quality breeding environments, but whose reproductive output is often influenced by stochastic events. Demographic estimates from this study are among the first for goldeneyes within Alaska. Life history patterns are known to vary geographically, therefore, we recommend further examination of life history patterns within the distribution of goldeneyes.     

Modulation of the adrenocortical response to acute stress with respect to brood value, reproductive success and survival in the Eurasian hoopoe

Reproducing parents face the difficult challenge of trading-off investment in current reproduction against presumed future survival and reproduction. Glucocorticoids are supposed to mediate this trade-off because the adrenocortical response to stress disrupts normal reproductive behaviour in favour of self-maintenance and own survival. According to the brood-value hypothesis, individuals with a low survival probability until the next reproductive season have to invest in current reproduction, a process driven by a down-regulation of their adrenocortical response. If the adrenocortical response to stress effectively mediates the trade-off between current reproduction versus future survival and reproduction, we expect a negative relationship with reproductive success and a positive correlation of the adrenocortical stress response with survival. We studied the relationship between corticosterone secretion in parents and their current brood value, reproductive success and survival in a short-lived multi-brooded bird, the Eurasian hoopoe Upupa epops. The adrenocortical response to acute handling stress was correlated with the brood value within the individual (first and second broods of the year) and between individuals. Birds breeding late in the season mounted a lower total corticosterone response to acute stress than birds breeding earlier, while females showed lower levels than males. We observed a negative relationship between the adrenocortical stress response and rearing success or fledging success in females, as predicted by the brood-value hypothesis. However, we could not evidence a clear link between the adrenocortical stress response and survival. Future research testing the brood-value hypothesis and trade-offs between current reproduction and future survival should also measure free corticosterone and carefully differentiate between cross-sectional (i.e. between-individual) and individual-based experimental studies.     

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.     

To breed or not to breed: past reproductive status and environmental cues drive current breeding decisions in a long-lived amphibian

Iteroparity is an adaptive response to uncertainty in reproductive success. However, spreading reproductive success over multiple reproduction events during a lifetime is constrained by adult mortality and the stochasticity associated with interactions between external factors and physiological states. The acquisition of information about environmental conditions during the growth of progeny and sufficient resources during the non-reproductive period are key factors for breeding success. Consequently, we hypothesized that long-lived animals may skip a breeding opportunity when information about unfavourable environmental conditions is available. In addition, nutritional constraints could prevent an animal from replenishing its reserves sufficiently to invest in the current breeding period. We investigated these questions using capture–recapture data from a 5-year study on a large population of yellow-bellied toads in a forest in north-eastern France. We took advantage of various advances in multi-state capture–recapture models (e.g. unobservable states and mixture models) to test our hypotheses. Our results show that the combined effects of rainfall deficit and the breeding/non-breeding state of individuals during the past breeding season affect breeding probability during the following breeding opportunity. We also found that females breed less frequently than males, suggesting that the overall energy cost of reproduction differs between genders. Finally, the results indicate that toad survival appears to be negatively influenced by rainfall deficits. We discuss the yellow-bellied toad’s reproductive behaviour in term of bet-hedging strategy and life history trait evolution.     

Drivers of age-specific survival in a long-lived seabird: contributions of observed and hidden sources of heterogeneity

1.?We assessed the relative influence of variability in recruitment age, dynamic reproductive investment (time-specific reproductive states) and frailty (unobserved differences in survival abilities across individuals) on survival in the black-legged kittiwake. Furthermore, we examined whether observed variability in survival trajectories was best explained by immediate reproductive investment, cumulative investment, or both. 2.?Individuals that delayed recruitment (≥ age 7) suffered a higher mortality risk than early recruits (age 3), especially later in life, suggesting that recruitment age may be an indicator of individual quality. Although recruitment age helped explain variation in survival, time-varying reproductive investment had a more substantial influence. 3.?The dichotomy of attempting to breed or not explained variability in survival across life better than other parameterizations of reproductive states such as clutch size, brood size or breeding success. In the kittiwake, the sinequanon condition to initiate reproduction is to hold a nest site, which is considered a very competitive activity. This might explain why attempting to breed is the key level of investment that affects survival, independent of the outcome (failure or success). 4.?Interestingly, the more individuals cumulate reproductive attempts over life, the lower their mortality risk, indicating that breeding experience may be a good indicator of parental quality as well. In contrast, attempting to breed at time t increased the risk of mortality between t and t + 1. We thus detected an immediate trade-off between attempting to breed and survival in this population; however, the earlier individuals recruited, and the more breeding experience they accumulated, the smaller the cost. 5.?Lastly, unobserved heterogeneity across individuals improved model fit more (1·3 times) than fixed and dynamic sources of observed heterogeneity in reproductive investment, demonstrating that it is critical to account for both sources of individual heterogeneity when studying survival trajectories. Only after simultaneously accounting for both sources of heterogeneity were we able to detect the 'cost' of immediate reproductive investment on survival and the 'benefit' of cumulative breeding attempts (experience), a proxy to individual quality.     

Individual heterogeneity in fitness in a long‐lived herbivore

Heterogeneity in the intrinsic quality and nutritional condition of individuals affects reproductive success and consequently fitness. Black brant (Branta bernicla nigricans) are long‐lived, migratory, specialist herbivores. Long migratory pathways and short summer breeding seasons constrain the time and energy available for reproduction, thus magnifying life‐history trade‐offs. These constraints, combined with long lifespans and trade‐offs between current and future reproductive value, provide a model system to examine the role of individual heterogeneity in driving life‐history strategies and individual heterogeneity in fitness. We used hierarchical Bayesian models to examine reproductive trade‐offs, modeling the relationships between within‐year measures of reproductive energy allocation and among‐year demographic rates of individual females breeding on the Yukon‐Kuskokwim Delta, Alaska, using capture–recapture and reproductive data from 1988 to 2014. We generally found that annual survival tended to be buffered against variation in reproductive investment, while breeding probability varied considerably over the range of clutch size‐laying date combinations. We provide evidence for relationships between breeding probability and clutch size, breeding probability and nest initiation date, and an interaction between clutch size and initiation date. Average lifetime clutch size also had a weak positive relationship with apparent survival probability. Our results support the use of demographic buffering strategies for black brant. These results also indirectly suggest associations among environmental conditions during growth, fitness, and energy allocation, highlighting the effects of early growth conditions on individual heterogeneity, and subsequently, lifetime reproductive investment.     

Dominance, reproduction and survival in banded mongooses: towards an egalitarian social system?

The banded mongoose, Mungos mungo, is a social species that forms multimale and multifemale family groups. Earlier studies suggest these family groups are relatively egalitarian with small differences in reproductive opportunities among individuals of different rank. In contrast, previous studies of other social mongooses have focused on species with more despotic control of reproduction (meerkats, Suricata suricatta, dwarf mongooses, Helogale parvula). In these species, the distribution of reproductive opportunities amongst individuals of different rank has met the predictions of reproductive skew theory: dominant individuals accrue greater reproductive benefits than subordinates, with subordinates breeding less often than dominants. In this paper we test how well two predictions of reproductive skew theory explain variance in measures of reproductive effort, and its correlates, in a wild population of banded mongooses in Queen Elizabeth National Park, Uganda. We measure dominance rank in males and females, and we investigate whether individuals of higher social rank accrue greater benefits than subordinates in terms of survival and reproduction. Banded mongoose dominance hierarchies showed linearity, but low reproductive skew. Rank was not significantly correlated with age. Furthermore, there were only small effects of dominance rank on nutritional levels, and no effects on reproduction and survival, suggesting that banded mongoose societies are indeed relatively egalitarian. No evidence of reproductive suppression was found and other forms of reproductive control were not observed. However, we do not exclude the possibility of increased reproductive competition in circumstances of higher ecological constraints. These findings show that reproductive skew theory is equally useful in explaining variation in reproduction in societies with low reproductive skew, as it is in explaining the allocation of reproductive effort in despotic social systems. Copyright 2001 The Association for the Study of Animal Behaviour.     

Immune challenge retards seasonal reproductive regression in rodents: evidence for terminal investment

Animals must balance investments in different physiological activities to allow them to maximize fitness in the environments they inhabit. These adjustments among reproduction, growth and survival are mandated because of the competing high costs of each process. Seasonally breeding rodents generally bias their investments towards reproduction when environmental conditions are benign, but shift these investments towards processes that promote survival, including immune activity, when environmental conditions deteriorate. Because survival probability of non-tropical small mammals is generally low in winter, under certain circumstances, these animals may not allocate resources to survival mechanisms in an effort to produce as many offspring as possible in the face of increased probability of death. Such 'terminal investments' have been described in passerines, but there are few examples of such phenomena in small mammals. Here, we show that male Siberian hamsters (Phodopus sungorus) challenged with lipopolysaccharide (a component of gram-negative bacteria that activates the immune system) induced a small, but significant, retardation of seasonal regression of the reproductive system relative to saline-injected hamsters. This delayed reproductive regression likely reflects a strategy to maintain reproductive function when survival prospects are compromised by infection.     

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.     

The impact of reproduction on gambian women: Does controlling for phenotypic quality reveal costs of reproduction?

Life history theory predicts that where resources are limited, investment in reproduction will cause a decline in body condition and ultimately may lower survival rates. We investigate the relationship between reproduction and mortality in women in rural Gambia. We use a number of different measures of reproductive investment: the timing of reproduction, intensity of reproduction, and cumulative reproductive investment (parity). Though giving birth is clearly a risk factor for increased mortality, we find limited evidence that the timing, intensity, or cumulative effects of reproduction have a survival cost. Instead, there is some evidence that women who have invested heavily in reproduction have higher survival than women with lower reproductive investment: both high parity and late age at last reproduction are associated with high survival. The only evidence for any cost of reproduction is that women who have given birth to twins (considered a marker of heavy investment in reproduction) have higher mortality rates than other women, after the age of 50 years. A potential confounding factor may be differences in health between women: particularly healthy women may be able to invest substantially in both reproduction and their own survival, leading to the positive correlations between survival and both parity and age at last birth we observe. To control for differences in health between women, we reanalyze the relationship between reproduction and mortality but include variables correlating with health in our models (height, BMI, and hemoglobin). Even when controlling for health, the positive correlation between investment in reproduction and survival remains unchanged.