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.     

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.     

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.     

Lifetime reproductive success in common buzzard, Buteo buteo: from individual variation to population demography

We studied annual and lifetime reproductive success (LRS) of both sexes of common buzzard Buteo buteo in eastern Westphalia, Germany. We followed a bottom‐up approach starting from individual breeding attempts, over lifetime reproductive success to derive population demography. Annual breeding performance and survival followed a quadratic relationship with breeding experience; individuals starting their breeding career were less likely to survive and breed successfully than birds of intermediate breeding experience. According to an analysis of selection gradients, both the opportunity and intensity of selection peaked in the early stages of the breeding career. The distribution of both LRS and another fitness measure, λ, was highly skewed, with ca 17% of adult birds producing 50% of fledglings in both sexes. Besides breeding life span and number of breeding attempts, habitat quality and plumage morph were significant predictors of LRS. There were strong differences in LRS and λ between the plumage morphs in both sexes: intermediate pigmented buzzards were much more successful than either dark or light ones. There was no significant difference between buzzard cohorts either in LRS or λ, nor did these fitness measures differ between individuals starting their breeding career at different conditions of food availability. Based on individual life histories, we formed a transition matrix and analysed its properties to study the population as a whole. This analysis showed that the population growth rate was close to unity (0.906, bootstrapped 95% confidence limits: 0.834 and 0.962). Analysis of reproductive values and elasticities further emphasised colour morph differences: the contribution of intermediate individuals to population growth greatly exceeded that of dark or light individuals. Thus most phenomena on all levels from individual breeding attempts over lifetime reproductive success to population demography can be explained by the fitness differences between the colour morphs with the intermediate morph maintaining the current population renewal potential.     

Longevity in cheetahs: the key to success?

An understanding of the factors governing reproductive success has fundamental implications for population demography, conservation, selection and adaptation. Although a consistent positive correlation between lifetime reproductive success and longevity has been reported for many iteroparous organisms, few studies have explored how longevity influences annual individual performance. In this study we show (1) that longevity and lifetime reproductive success are positively but not linearly correlated, (2) that short-lived individuals have higher annual reproductive success, (3) that the generally lower success of the last breeding occasion increased with females' longevity, and (4) that long-lived females have higher chances of rearing long-lived females. We suggest that experience and the increase in the number of reproductive events with longevity are key processes leading to a strong correlation between (1) lifetime reproductive success and longevity and (2) mother and daughter longevities. Our results demonstrate the importance of long term studies that follow multiple generations in gaining a full understanding of the factors affecting reproductive success.     

Fitness and behavioral correlates of pre-stress and stress-induced plasma cortisol titers in pink salmon (Oncorhynchus gorbuscha) upon arrival at spawning grounds

Semelparous Pacific salmon (Onchorynchus spp.) serve as an excellent model for examining the relationships between life history, behavior and individual variation in glucocorticoid (GC) stress hormone levels because reproductive behaviors are highly variable between individuals and failure to reproduce results in zero fitness. Pink salmon (O. gorbuscha) were intercepted upon arrival at the spawning grounds across three time periods. Pre-stress and stress-induced plasma cortisol concentrations were assessed in relation to behavior, longevity and reproductive success. Results revealed differences between sexes and with arrival time. The study period marked a year of high reproductive success and only nine females (12% of sample) failed to spawn. Female pre-spawn mortalities were characterized by significantly elevated stress-induced cortisol concentrations and decreased longevity as well as pre-stress cortisol above the normal range in pink salmon from the study area. Interestingly, reproductive behaviors were only associated with pre-stress cortisol levels. For females, aggression and mate interaction time were reduced in individuals with elevated pre-stress cortisol concentrations. In males, a similar negative relationship between pre-stress cortisol concentration and mate interaction time was detected. The observed behavioral correlations are likely a factor of social status where dominant individuals, known to have higher reproductive success, are characterized by lower cortisol levels relative to subordinate conspecifics. Findings show both elevated pre-stress and stress-induced cortisol concentrations at arrival to the spawning grounds to be associated with reduced survival.     

Fitness and behavioral correlates of pre-stress and stress-induced plasma cortisol titers in pink salmon (Oncorhynchus gorbuscha) upon arrival at spawning grounds

Semelparous Pacific salmon (Onchorynchus spp.) serve as an excellent model for examining the relationships between life history, behavior and individual variation in glucocorticoid (GC) stress hormone levels because reproductive behaviors are highly variable between individuals and failure to reproduce results in zero fitness. Pink salmon (O. gorbuscha) were intercepted upon arrival at the spawning grounds across three time periods. Pre-stress and stress-induced plasma cortisol concentrations were assessed in relation to behavior, longevity and reproductive success. Results revealed differences between sexes and with arrival time. The study period marked a year of high reproductive success and only nine females (12% of sample) failed to spawn. Female pre-spawn mortalities were characterized by significantly elevated stress-induced cortisol concentrations and decreased longevity as well as pre-stress cortisol above the normal range in pink salmon from the study area. Interestingly, reproductive behaviors were only associated with pre-stress cortisol levels. For females, aggression and mate interaction time were reduced in individuals with elevated pre-stress cortisol concentrations. In males, a similar negative relationship between pre-stress cortisol concentration and mate interaction time was detected. The observed behavioral correlations are likely a factor of social status where dominant individuals, known to have higher reproductive success, are characterized by lower cortisol levels relative to subordinate conspecifics. Findings show both elevated pre-stress and stress-induced cortisol concentrations at arrival to the spawning grounds to be associated with reduced survival.     

Body Condition Indices Predict Reproductive Success but Not Survival in a Sedentary,Tropical Bird

Body condition may predict individual fitness because those in better condition have more resources to allocate towards improving their fitness. However, the hypothesis that condition indices are meaningful proxies for fitness has been questioned. Here, we ask if intraspecific variation in condition indices predicts annual reproductive success and survival. We monitored a population of Neochmia phaeton (crimson finch), a sedentary, tropical passerine, for reproductive success and survival over four breeding seasons, and sampled them for commonly used condition indices: mass adjusted for body size, muscle and fat scores, packed cell volume, hemoglobin concentration, total plasma protein, and heterophil to lymphocyte ratio. Our study population is well suited for this research because individuals forage in common areas and do not hold territories such that variation in condition between individuals is not confounded by differences in habitat quality. Furthermore, we controlled for factors that are known to impact condition indices in our study population (e.g., breeding stage) such that we assessed individual condition relative to others in the same context. Condition indices that reflect energy reserves predicted both the probability of an individual fledging young and the number of young produced that survived to independence, but only during some years. Those that were relatively heavy for their body size produced about three times more independent young compared to light individuals. That energy reserves are a meaningful predictor of reproductive success in a sedentary passerine supports the idea that energy reserves are at least sometimes predictors of fitness. However, hematological indices failed to predict reproductive success and none of the indices predicted survival. Therefore, some but not all condition indices may be informative, but because we found that most indices did not predict any component of fitness, we question the ubiquitous interpretation of condition indices as surrogates for individual quality and fitness.     

Determinants of quality in a long-lived colonial species

1. In many animal populations a small proportion of individuals produce the majority of surviving offspring, but the underlying mechanisms are unclear. Behaviour may be an important determinant of variation in fitness: 'high-quality' individuals may have enhanced abilities in foraging or predator and parasite avoidance. 2. The role of behaviour in determining variation in quality was examined using the common guillemot Uria aalge, a monogamous seabird with biparental care. Using a novel mixed model approach, we analysed binary data on breeding success of each pair attempting to breed in each year with variables critical to breeding success (timing of breeding; inferred age; breeding experience and success; number of nest sites and partners) as fixed effects. Random effects for year, male, female and each distinct pairing of a male and a female were included in the model, allowing a quality estimate to be derived for each individual and pair. A range of behaviours associated with breeding were examined in relation to these quality estimates. 3. Breeding success declined with timing of breeding, and increased initially with age before declining in old age. It increased with previous successful experience, not breeding experience per se, until senescence effects became apparent. For males, breeding success declined with increasing numbers of mates. 4. The most important behavioural determinants of quality operated at the level of the pair, with the time mates spent together at the site and chick feeding rates both positively related to quality. At the individual level, trip durations and feeding rates were associated with female but not male quality, suggesting that pair quality was operating principally through the female. However, removal of laying date, the most important component in the binomial model, confirmed that the pair effect was much larger than the female effect. 5. This study demonstrates the potential of mixed modelling to determine quality estimates based on long-term breeding histories. The probability of a successful reproductive attempt was explained by the timing of breeding, age, successful breeding experience and number of mates. Behaviour was an important proximate mechanism underlying quality, in particular the foraging abilities of the pair, and the female's contribution to offspring provisioning. In species with biparental care, behavioural correlates of quality operate most strongly at the scale of the breeding pair, because contributions from both individuals are required for a successful outcome.     

Lifetime offspring production in relation to breeding lifespan, attractiveness, and mating status in male collared flycatchers

As a comprehensive fitness parameter, lifetime reproductive success (LRS) is influenced by many different environmental and genetic factors, among which longevity is one of the most important. These factors can be reflected in secondary sexual characters, which may affect the life histories of individuals via social relations with conspecifics. Facultative polygyny in birds is another conspicuous reproductive trait that potentially increases male reproductive success, but lifetime success data in relation to polygyny are scarce. Here, we used 17?years of breeding data to quantify the LRS of male collared flycatchers (Ficedula albicollis) on the basis of lifetime recruitment of offspring. Breeding lifespan showed a positive relationship with LRS, and it was also significantly associated with mean recruitment of offspring per breeding year. Body size and sexually selected forehead patch size did not predict the number of recruits. Polygyny was positively associated with LRS, but when we corrected for lifespan, this relationship disappeared. Our results demonstrate that the relationship between longevity and LRS is not explained by the higher number of reproductive attempts when living longer, and question the adaptive value of polygyny in this population. The lack of association between forehead patch size and recruitment suggests that forehead patch is a poor indicator of phenotypic quality in our birds.     

Long‐lived and heavier females give birth earlier in roe deer

In seasonal environments, parturition of most vertebrates generally occurs within a short time‐window each year. This synchrony is generally interpreted as being adaptive, as early born young survive better over the critical season than late born young. Among large herbivores, the factors involved in driving among‐ and within‐individual variation in parturition date are poorly understood. We explored this question by analyzing the relative importance of attributes linked to female quality (longevity, median adult body mass and cohort), time‐dependent attributes linked to female condition (reproductive success the previous year, relative annual body mass and offspring cohort (year)), and age in shaping observed variation in parturition date of roe deer. A measure of quality combining the effects of female longevity and median adult body mass accounted for 11% of the observed among‐individual variation in parturition date. Females of 2 yr old give birth 5 d later than older females. Our study demonstrates that high quality (heavy and long‐lived) females give birth earlier than low quality females. Temporally variable attributes linked to female condition, such as reproductive success in the previous year and relative annual body mass, had no detectable influence on parturition date. We conclude that parturition date, a crucial determinant of reproductive success, is shaped by attributes linked to female quality rather than by time‐dependent attributes linked to female condition in income breeders (individuals that rely on current resource intake rather than on accumulated body reserves to offset the increased energy requirements due to reproduction) such as roe deer.     

HIGHER FITNESS FOR PHILOPATRIC THAN FOR IMMIGRANT MALES IN A SEMI-ISOLATED POPULATION OF GREAT REED WARBLERS

To compare the fitness of philopatric and immigrant individuals we examined the lifetime reproductive success of 116 male and 137 female great reed warblers. The study was carried out in a semi-isolated population in Sweden and covered breeding adults hatched between 1985 and 1993. Lifetime fitness, measured as life time number of fledglings and offspring recruits, was lower for immigrant than for philopatric males. We found no such relationships for females. The difference in reproductive success could not be explained by immigrant males having lower phenotypic quality because they had similar life span, spring arrival date, and territory quality as philopatric males. The lower lifetime fitness among immigrant than philopatric males appeared to result from reduced mating success. This suggests that females are reluctant to mate with immigrant males despite their apparently similar phenotypic quality. Though it is not known whether females gain in fitness by avoiding matings with immigrant males, it is notable that immigrant males have smaller song repertoires than philopatric males. Large repertoires, previously shown to sexually arouse great reed warbler females, correlate with the occurrence of extrapair paternity and postfledging survival of offspring in our population.     

Age‐dependent changes in infidelity in Seychelles warblers

Extra‐pair paternity (EPP) is often linked to male age in socially monogamous vertebrates; that is, older males are more likely to gain EPP and less likely to be cuckolded. However, whether this occurs because males improve at gaining paternity as they grow older, or because “higher quality” males that live longer are preferred by females, has rarely been tested, despite being central to our understanding of the evolutionary drivers of female infidelity. Moreover, how extra‐pair reproduction changes with age within females has received even less attention. Using 18 years of longitudinal data from an individually marked population of Seychelles warblers (Acrocephalus sechellensis), we found considerable within‐individual changes in extra‐pair reproduction in both sexes: an early‐life increase and a late‐life decline. Furthermore, males were cuckolded less as they aged. Our results indicate that in this species age‐related patterns of extra‐pair reproduction are determined by within‐individual changes with age, rather than differences among individuals in longevity. These results challenge the hypothesis—based on longevity reflecting intrinsic quality—that the association between male age and EPP is due to females seeking high‐quality paternal genes for offspring. Importantly, EPP accounted for up to half of male reproductive success, emphasizing the male fitness benefits of this reproductive strategy. Finally, the occurrence of post‐peak declines in extra‐pair reproduction provides explicit evidence of senescence in infidelity in both males and females.     

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.     

Adaptive female choice for middle-aged mates in a lekking sandfly

Most theoretical models of age-related mate choice predict that females should prefer older males because they have proven survival ability. An alternative view is that older males represent inferior mates because of negative genetic correlations between early and late fitness components, or because older males have traded off longevity against other fitness components, have accumulated deleterious germ-line mutations, or are less well adapted to current conditions than more recently born individuals. While numerous studies have reported female choice for older males, few have explicitly examined the fitness consequences of such a preference. We present evidence from a lekking sandfly, Lutzomyia longipalpis, showing that choosy females discriminate against older males and gain a fitness benefit from their choice. When permitted free choice from an aggregation consisting of males aged zero to two days (young), four to six days (middle-aged) and eight to ten days (old), females preferentially mated with middle-aged males, but all measures of female reproductive success were independent of male age. In contrast, when a second set of females was randomly assigned single virgin males of known age, the eggs of those paired to old mates exhibited lower hatching success than the eggs of females mated to young or middle-aged males. These results suggest that females avoid mating with older males because they represent poorer quality mates. Age-related differences in male quality may have a genetic basis, but could equally well arise through a phenotypic decline in sperm quality or sperm transfer ability with male age. The lack of evidence of female discrimination against older males from other studies may be because these did not explore the reproductive success of the full age range of males.     

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.     

Live fast, die young: trade-offs between fitness components and sexually antagonistic selection on weaponry in Soay sheep

Males are predicted to compete for reproductive opportunities, with sexual selection driving the evolution of large body size and weaponry through the advantage they confer for access to females. Few studies have explored potential trade-offs of investment in secondary sexual traits between different components of fitness or tested for sexually antagonistic selection pressures. These factors may provide explanations for observed polymorphisms in both form and quality of secondary sexual traits. We report here an analysis of selection on horn phenotype in a feral population of Soay sheep (Ovis aries) on the island of Hirta, St. Kilda, Scotland. Soay sheep display a phenotypic polymorphism for horn type with males growing either normal or reduced (scurred) horns, and females growing either normal, scurred, or no (polled) horns; further variation in size exists within horn morphs. We show that horn phenotype and the size of the trait displayed is subject to different selection pressures in males and females, generating sexually antagonistic selection. Furthermore, there was evidence of a trade-off between breeding success and longevity in normal-horned males, with both the normal horn type and larger horn size being associated with greater annual breeding success but reduced longevity. Therefore, selection through lifetime breeding success was not found to act upon horn phenotype in males. In females, a negative association of annual breeding success within the normal-horned phenotype did not result in a significant difference in lifetime fitness when compared to scurred individuals, as no significant difference in longevity was found. However, increased horn size within this group was negatively associated with breeding success and longevity. Females without horns (polled) suffered reduced longevity and thus reduced lifetime breeding success relative the other horn morphs. Our results therefore suggest that trade-offs between different components of fitness and antagonistic selection between the sexes may maintain genetic variation for secondary sexual traits within a population.     

Senescence and carryover effects of reproductive performance influence migration,condition, and breeding propensity in a small shorebird

Breeding propensity, the probability that an animal will attempt to breed each year, is perhaps the least understood demographic process influencing annual fecundity. Breeding propensity is ecologically complex, as associations among a variety of intrinsic and extrinsic factors may interact to affect an animal's breeding decisions. Individuals that opt not to breed can be more difficult to detect than breeders, which can (1) lead to difficulty in estimation of breeding propensity, and (2) bias other demographic parameters. We studied the effects of sex, age, and population reproductive success on the survival and breeding propensity of a migratory shorebird, the piping plover (Charadrius melodus ), nesting on the Missouri River. We used a robust design Barker model to estimate true survival and breeding propensity and found survival decreased as birds aged and did so more quickly for males than females. Monthly survival during the breeding season was lower than during migration or the nonbreeding season. Males were less likely to skip breeding (range: 1–17%) than females (range: 3–26%; β_sex = ?0.21, 95% CI: ?0.38 to ?0.21), and both sexes were less likely to return to the breeding grounds following a year of high reproductive success. Birds that returned in a year following relatively high population‐wide reproductive output were in poorer condition than following a year with lower reproductive output. Younger adult birds and females were more likely to migrate from the breeding area earlier than older birds and males; however, all birds stayed on the breeding grounds longer when nest survival was low, presumably because of renesting attempts. Piping plovers used a variety of environmental and demographic cues to inform their reproduction, employing strategies that could maximize fitness on average. Our results support the “disposable soma” theory of aging and follow with predictions from life history theory, exhibiting the intimate connections among the core ecological concepts of senescence, carryover effects, and life history.     

Delayed timing of breeding as a cost of reproduction

Timing of breeding is a trait with considerable individual variation, often closely linked to fitness because of seasonal declines in reproduction. The drivers of this variation have received much attention, but how reproductive costs may influence the timing of subsequent breeding has been largely unexplored. We examined a population of northern wheatears Oenanthe oenanthe to compare three groups of individuals that differed in their timing of breeding termination and reproductive effort to investigate how these factors may carry over to influence reproductive timing and reproductive output in the following season. Compared to females that bred successfully, females that put in less effort and terminated breeding early due to nest failure tended to arrive and breed earlier in year 2 (mean advancement = 2.2 and 3.3 d respectively). Females that spent potentially more effort and terminated breeding later due to production of a replacement clutch after nest failure, arrived later than other females in year 2. Reproductive output (number of fledglings) in year 2 differed between the three groups as a result of group‐level differences in the timing of breeding in combination with the general seasonal decline in reproductive output. Our study shows that the main cost of reproduction was apparent in the timing of arrival and breeding in this migratory species. Hence, reproductive costs can arise through altered timing of breeding since future reproductive success (including adult survival) is often dependent on the timing of breeding in seasonal systems.     

Time spent in distinct life history stages has sex‐specific effects on reproductive fitness in wild Atlantic salmon

In species with complex life cycles, life history theory predicts that fitness is affected by conditions encountered in previous life history stages. Here, we use a 4‐year pedigree to investigate if time spent in two distinct life history stages has sex‐specific reproductive fitness consequences in anadromous Atlantic salmon (Salmo salar). We determined the amount of years spent in fresh water as juveniles (freshwater age, FW, measured in years), and years spent in the marine environment as adults (sea age, SW, measured in sea winters) on 264 sexually mature adults collected on a river spawning ground. We then estimated reproductive fitness as the number of offspring (reproductive success) and the number of mates (mating success) using genetic parentage analysis (>5,000 offspring). Sea age is significantly and positively correlated with reproductive and mating success of both sexes whereby older and larger individuals gained the highest reproductive fitness benefits (females: 62.2% increase in offspring/SW and 34.8% increase in mate number/SW; males: 201.9% offspring/SW and 60.3% mates/SW). Younger freshwater age was significantly related to older sea age and thus increased reproductive fitness, but only among females (females: ?33.9% offspring/FW and ?32.4% mates/FW). This result implies that females can obtain higher reproductive fitness by transitioning to the marine environment earlier. In contrast, male mating and reproductive success was unaffected by freshwater age and more males returned at a younger age than females despite the reproductive fitness advantage of later sea age maturation. Our results show that the timing of transitions between juvenile and adult phases has a sex‐specific consequence on female reproductive fitness, demonstrating a life history trade‐off between maturation and reproduction in wild Atlantic salmon.