Age-specific resource investment strategies: evidence from female Richardson's ground squirrels (Spermophilus richardsonii)

To avoid a possible cost to their future survival and/or reproduction, individuals must balance their somatic and reproductive investments. The van Noordwijk and De Jong model of resource investment predicts that investments into reproduction and soma can vary among individuals of a population based on the variation in the total amount of energy that individuals acquire. With principal components analysis (PCA), we created two axes of life history for female Richardson's ground squirrels Spermophilus richardsonii : an index of total energy investment (PC1) and an index of investment tactic (PC2). Using these indices, we examined patterns of resource allocation to reproductive and somatic investments. Because yearling female Richardson's ground squirrels complete growth to adult size during pregnancy and early lactation, their somatic needs exceed those of older, fully grown females. Therefore, we predicted that yearlings would show more evidence of a tradeoff between reproductive and somatic investments compared with older females. Both yearling and older females invested four to five times more mass into their litters than into their own body mass. With increasing total investment, yearling females increased investment in both reproduction and themselves, whereas older females invested relatively more in reproduction than themselves. Regardless of age, females that emerged heavier from hibernation invested fewer resources into themselves and more into their litters. Variation in total energy investment and investment tactic indices was similar for yearling and older females. Contrary to our prediction, however, yearling females showed positive associations between reproductive and somatic investments, whereas older females exhibited showed no significant association between reproductive and somatic investments.     

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.     

Age-specific variation in survival, reproductive success and offspring quality in red squirrels: evidence of senescence

Individual performance is expected to decrease with age because of senescence. We analyzed long-term data collected on a North American red squirrel population to assess the influence of age on body mass, survival and reproductive performance, and to study the effects of sex and of environmental conditions during early life on senescence patterns. Mass of males and females did not decrease at the end of life, possibly because body mass mostly reflects overall size in income breeders such as red squirrels. On the other hand, we found evidence of senescence in survival of both sexes and, to a lesser extent, in female reproductive traits. When compared to females, males had both higher survival and delayed decrease in survival, suggesting a weaker senescence in males. The offspring survival from weaning to one year of age also decreased with increasing mother age. This suggests that older females produce juveniles of lower quality, providing evidence of an intergenerational effect of mother's age on juveniles' fitness. Finally, our results indicate that variations in food conditions during early life influenced the reproductive tactics of females in the first years of their life, but not senescence patterns.     

Reproductive effort during gestation and lactation by Richardson's ground squirrels

Summary Reproductive effort by yearling and older female Richardson's ground squirrels was studied over a 4-year period in southern Alberta by obtaining serial weight records from marked individuals to compare the mother's mass at critical points in the annual cycle (emergence from hibernation, estrus, parturition, and litter emergence) with her litter's mass at birth and weaning. Yearlings weighed only 80% of older adults at emergence from hibernation, but they mated at the same time as older females, attained adult mass coincident with pregnancy, and weaned litters that were not significantly smaller in size or mass than those of older females. Age and maternal mass were weak predictors of litter size and litter mass. Of the net increase in mass of the combined mother-litter unit during gestation, over half (60% of 139 g for yearlings; 52% of 127 g for older females) was attributable to an increase in the mother's own mass, whereas during lactation almost all of the net increase (93% of 545 g for yearlings; 96% of 567 g for older females) was attributable to an increase in the litter's mass. On a daily basis, deposition of mass in the litter was 6 times greater during lactation than gestation. On average, neonates weighed 2.3% (6.5 g) of maternal mass at birth and 23.1% (81 g) at emergence from the natal burrow; offspring masses at birth and at emergence were significantly negatively correlated with litter size. On average, litters weighed 16.3% (48 g) of maternal mass at birth and 157.5% (578 g) at emergence from the natal burrow. Compared with other hibernating sciurids, Richardson's ground squirrels have a similar offspring mass relative to maternal mass both at birth and at emergence from the natal burrow. However, because of the large litter size (typically 6–8), absolute reproductive effort, measured either as litter mass at birth or at natal emergence, is large for the body size of the species.     

Terminal investment and senescence in rhesus macaques (Macaca mulatta) on Cayo Santiago

Long-lived iteroparous species often show aging-related changes in reproduction that may be explained by 2 non-mutually exclusive hypotheses. The terminal investment hypothesis predicts increased female reproductive effort toward the end of the life span, as individuals have little to gain by reserving effort for the future. The senescence hypothesis predicts decreased female reproductive output toward the end of the life span due to an age-related decline in body condition. Nonhuman primates are ideal organisms for testing these hypotheses, as they are long lived and produce altricial offspring heavily dependent on maternal investment. In this study, we integrated 50 years of continuous demographic records for the Cayo Santiago rhesus macaque (Macaca mulatta) population with new morphometric and behavioral data to test the senescence and terminal investment hypotheses. We examined relationships between maternal age and activity, mother and infant body condition, interbirth intervals, measures of behavioral investment in offspring, and offspring survival and fitness to test for age-associated declines in reproduction that would indicate senescence, and for age-associated increases in maternal effort that would indicate terminal investment. Compared with younger mothers, older mothers had lower body mass indices and were less active, had longer interbirth intervals, and spent more time in contact with infants, but had infants of lower masses and survival rates. Taken together, our results provide strong evidence for the occurrence of reproductive senescence in free-ranging female rhesus macaques but are also consistent with some of the predictions of the terminal investment hypothesis.     

Size delays female senescence in a medium sized marsupial: The effects of maternal traits on annual fecundity in the northern brown bandicoot (Isoodon macrourus)

The degree to which females allocate resources between current reproduction, future fecundity and survival is a central theme in life history theory. We investigated two hypotheses proposed to explain patterns of reproductive investment, terminal investment and senescence, by examining the effects of maternal traits (age and maternal mass) on annual fecundity in female northern brown bandicoots, Isoodon macrourus (Marsupialia: Peramelidae). We found that annual fecundity in females declined in their final year of reproduction, indicating reproductive senescence. Maternal mass significantly influenced the rate of senescence and, in turn, a female's lifetime reproductive output. Mass had little effect on fecundity in 1st and 2nd year females, but a positive relationship with fecundity in 3rd year females. This meant that heavy, 3rd year females did not suffer the decline in fecundity shown in light 3rd year females. For 1st year females, mass and leg length increased between their first and second reproductive seasons, indicating a temporary shift, from the allocation of resources to reproduction, to increasing condition or structural size post their first breeding event. There were no net changes to body mass in subsequent years. We suggest that this year of post‐reproductive growth has important consequences for senescent effects on reproduction. Overall, results provided support for the effects of senescence on annual fecundity. Our findings were not consistent with the terminal investment hypothesis; reproductive output did not increase in females' final reproductive season despite a rapid decline in survival. However, this notion cannot be entirely dismissed; other measures of reproductive performance not examined here (e.g. offspring mass) may have provided an indication that females did increase their effort at the end of their lifespan. This study highlights the difficulty of measuring reproductive costs and the importance of understanding the combined effects of specific characteristics of an individual when interpreting reproductive strategies in iteroparous organisms.     

Contrasting life histories in neighbouring populations of a large mammal

Background

A fundamental life history question is how individuals should allocate resources to reproduction optimally over time (reproductive allocation). The reproductive restraint hypothesis predicts that reproductive effort (RE; the allocation of resources to current reproduction) should peak at prime-age, whilst the terminal investment hypothesis predicts that individuals should continue to invest more resources in reproduction throughout life, owing to an ever-decreasing residual reproductive value. There is evidence supporting both hypotheses in the scientific literature.

Methodology/Principal Findings

We used an uncommonly large, 38 year dataset on Alpine chamois (Rupicapra rupicapra) shot at various times during the rutting period to test these two hypotheses. We assumed that body mass loss in rutting males was strongly related to RE and, using a process-based approach, modelled how male relative mass loss rates varied with age. For different regions of our study area, we provide evidence consistent with different hypotheses for reproductive allocation. In sites where RE declined in older age, this appears to be strongly linked to declining body condition in old males. In this species, terminal investment may only occur in areas with lower rates of body mass senescence.

Conclusions/Significance

Our results show that patterns of reproductive allocation may be more plastic than previously thought. It appears that there is a continuum from downturns in RE at old age to terminal investment that can be manifest, even across adjacent populations. Our work identifies uncertainty in the relationship between reproductive restraint and a lack of competitive ability in older life (driven by body mass senescence); both could explain a decline in RE in old age and may be hard to disentangle in empirical data. We discuss a number of environmental and anthropogenic factors which could influence reproductive life histories, underlining that life history patterns should not be generalised across different populations.     

Age‐specific reproduction and disposable soma in an urban population of Common Blackbirds Turdus merula

The mechanism of senescence is an important subject of current research, but our knowledge of the factors influencing the rate of ageing in naturally occurring populations remains rudimentary. Evolutionary theories of senescence predict that investment in reproduction in early life should come at the cost of reduced somatic maintenance and thus result in earlier or more rapid senescence. We use data on the complete reproductive histories of 431 Common Blackbirds (222 males and 209 females) collected during a 19‐year study of the ecology of an urban population of this species to test the main hypotheses addressing the issue of senescence. On average, the birds in this population survived for 3.7 (± 1.9 sd) years. Reproductive success in females peaked at the age of 4, but in males remained stable until the 5th year of life. We observed declines in reproductive success, indicative of senescence, after the peak years in both sexes. The mechanism of age‐related changes in the reproduction of females confirms the individual improvement and selective disappearance hypotheses. In the case of males, the increase in reproductive performance comes as a consequence of the disappearance of poor reproducers. The parental investment associated with early life fecundity (the first two breeding seasons in males and females) impairs the breeding success of females later on. Contrary to expectations, there was no negative impact of high early life fecundity on either mortality or lifespan. Individuals of both sexes with a high early life fecundity had a higher lifetime reproductive success than those in which early life fecundity was low. Hence, the most profitable strategy is to maximize reproductive effort in the early stages of life. This yields the highest lifetime reproductive success, despite the increased impact of senescence, especially in females. These results are consistent with the disposable soma hypothesis.     

The "hibernation induction trigger": specificity and validity of bioassay using the 13-lined ground squirrel

Even though the existence of the blood-borne "hibernation induction trigger" has been reported in the 13-lined ground squirrel, transfusion of plasma from hibernating rodents with other hibernating species as the recipients failed to induce the occurrence of summer hibernation. In order to verify whether the response to the "trigger" substance is species specific, the present study was carried out to compare the effect of plasma from hibernating Richardson's ground squirrels on the incidence of summer hibernation in both juvenile Richardson's and adult 13-lined ground squirrels. In two series of experiments, 13-lined ground squirrels entered hibernation quite readily independent of the treatment. The rate of occurrence of hibernation ranged from 78% after sham injection to 86% after warm saline, fresh summer active plasma, and fresh hibernating plasma, respectively. There were no differences in the number of hibernation bouts and the number of days in hibernation after each treatment. In contrast, none of the juvenile Richardson's ground squirrels entered hibernation after any of the treatments up to the end of the 8-week observation period. These results not only argue against the existence of blood-borne "trigger" substance, at least in the Richardson's ground squirrel, but also caution against the use of the 13-lined ground squirrel as a standard test animal for the bioassay of the "trigger" substance.     

Reproductive Decisions in Female European Ground Squirrels: Factors Affecting Reproductive Output and Maternal Investment

Physiological and behavioural parameters associated with reproductive effort and success were investigated in female European ground squirrels Spermophilus citellus . The proportion of reproductive (lactating) females in the study population was over 90% and was not related to age. Timing of oestrus and ovulation was found to be affected by the female's emergence date and condition. Females with low emergence mass showed delayed oestrus. Differences in ovulation dates were shown to affect reproductive output in terms of litter size and sex ratio. Early litters were larger and male biased. X-ray techniques were used to determine intrauterine litter size in individual females. The results indicated that litter size and sex ratio were fixed prenatally. Lactation costs were reflected in the intensity of mass loss and duration of lactation. Mass loss varied with litter size, in that females with large litters showed a more rapid loss than others. The second parental investment parameter, lactation duration, varied among individual females and was dependent on the timing of reproduction and litter size (except yearlings). Early born litters, which were, in most cases, larger than later ones, were nursed longer. Prolonged lactation periods affected female condition in that they started prehibernation fattening later and entered hibernation with a lower mass than individuals that had shorter lactation periods. Yearling females probably could not afford the energetic costs of long lactation, independent of their offspring number. These results indicated that females with higher reproductive output and higher investment were unable to compensate these costs before hibernation. Consequences for these individuals could therefore be lower over-winter survival or a delayed oestrus in the following season.     

Increasing returns in the life history of Columbian ground squirrels

1. We examined positive associations and trade-offs of maternal and reproductive traits in a population of Columbian ground squirrels, Spermophilus columbianus .
2. Structural size, body condition, mother's personal allocation to body mass during reproduction, and timing of littering were estimated for live-trapped reproductive females that were observed during an 8-year period, and were compared to litter mass, litter size, and average pup mass using path analyses.
3. Mothers exhibited age-structured traits that influenced reproductive patterns. Yearling mothers were significantly smaller, bred later, and had smaller litters than older females. Mothers that gained more body mass during reproduction and older mothers in good body condition that were structurally large had larger litters.
4. Early seasonal timing of littering was an important positive influence on successful reproduction by older mothers only in early breeding seasons and in years when conditions for reproduction were good for all females.
5. The number of offspring that survived to 1 year of age was most strongly associated with litter mass and litter size; date of breeding was of secondary influence, with earlier litters exhibiting greater success.
6. In general, mothers that gained the most in body mass during reproduction were concurrently more successful in weaning large litters (perhaps due to better quality of foraging habitat).
7. In addition to expected reproductive trade-offs, reproduction by Columbian ground squirrels exhibited positive associations of life-history traits that may reflect evolutionary increasing returns.     

Are reproductive and somatic senescence coupled in humans? Late, but not early, reproduction correlated with longevity in historical Sami women

Evolutionary theory of senescence emphasizes the importance of intense selection on early reproduction owing to the declining force of natural selection with age that constrains lifespan. In humans, recent studies have, however, suggested that late-life mortality might be more closely related to late rather than early reproduction, although the role of late reproduction on fitness remains unclear. We examined the association between early and late reproduction with longevity in historical post-reproductive Sami women. We also estimated the strength of natural selection on early and late reproduction using path analysis, and the effect of reproductive timing on offspring survival to adulthood and maternal risk of dying at childbirth. We found that natural selection favoured both earlier start and later cessation of reproduction, and higher total fecundity. Maternal age at childbirth was not related to offspring or maternal survival. Interestingly, females who produced their last offspring at advanced age also lived longest, while age at first reproduction and total fecundity were unrelated to female longevity. Our results thus suggest that reproductive and somatic senescence may have been coupled in these human populations, and that selection could have favoured late reproduction. We discuss alternative hypotheses for the mechanisms which might have promoted the association between late reproduction and longevity.     

Testing the reproductive and somatic trade‐off in female Columbian ground squirrels

Energetic trade‐offs in resource allocation form the basis of life‐history theory, which predicts that reproductive allocation in a given season should negatively affect future reproduction or individual survival. We examined how allocation of resources differed between successful and unsuccessful breeding female Columbian ground squirrels to discern any effects of resource allocation on reproductive and somatic efforts. We compared the survival rates, subsequent reprodction, and mass gain of successful breeders (females that successfully weaned young) and unsuccessful breeders (females that failed to give birth or wean young) and investigated “carryover” effects to the next year. Starting capital was an important factor influencing whether successful reproduction was initiated or not, as females with the lowest spring emergence masses did not give birth to a litter in that year. Females that were successful and unsuccessful at breeding in one year, however, were equally likely to be successful breeders in the next year and at very similar litter sizes. Although successful and unsuccessful breeding females showed no difference in over winter survival, females that failed to wean a litter gained additional mass during the season when they failed. The next year, those females had increased energy “capital” in the spring, leading to larger litter sizes. Columbian ground squirrels appear to act as income breeders that also rely on stored capital to increase their propensity for future reproduction. Failed breeders in one year “prepare” for future reproduction by accumulating additional mass, which is “carried over” to the subsequent reproductive season.     

Lipid synthesis and deposition by adult Richardson's ground squirrels in the natural environment

Summary Adult male Richardson's ground squirrels,Spermophilus richardsonii, were estimated to have emerged from hibernation in late February to early March, and adult females in mid to late March. Half of the females trapped in late March were not pregnant, as against 10% after that time. In late March males and all females had similar WAT (white adipose tissue) deposits. Between late March and early June, WAT deposits in males increased from 14 g to 64 g (a rate of 5.6 g per week). In non-parous females WAT deposits increased from 13 g to 48 g from late March to late May (4.2 g per week). Fat deposits decreased during lactation but thereafter increased from 8 g to 29 g (a rate of 6.0 g per week) between early May and early June. In males the rate of fatty acid synthesis in BAT (brown adipose tissue), liver and WAT did not change from late March to late May, and rates in the corresponding tissues of non-pregnant females were similar to those in males. Fatty acid synthesis decreased during late pregnancy and lactation. After lactation, the rate of fatty acid synthesis in all tissues increased to that in males and non-pregnant females. Males initiated fattening 5–7 weeks earlier than females. It is concluded that compared with adult males, the later immergence of adult female Richardson's ground squirrels into hibernation is due primarily to later initiation of fattening and less to differences in rate of lipid synthesis after the reproductive period. Rates of fatty acid synthesis in liver and BAT were several times greater than that in WAT. The former tissues may contribute fatty acids for prehibernatory fattening.Abbreviations BAT brown adipose tissue - WAT white adipose tissue     

Survival costs of reproduction predict age-dependent variation in maternal investment

Life-history theory predicts that older females will increase reproductive effort through increased fecundity. Unless offspring survival is density dependent or female size constrains offspring size, theory does not predict variation in offspring size. However, empirical data suggest that females of differing age or condition produce offspring of different sizes. We used a dynamic state-variable model to determine when variable offspring sizes can be explained by an interaction between female age, female state and survival costs of reproduction. We found that when costs depend on fecundity, young females with surplus state increase offspring size and reduce number to minimize fitness penalties. When costs depend on total reproductive effort, only older females increase offspring size. Young females produce small offspring, because decreasing offspring size is less expensive than number, as fitness from offspring investment is nonlinear. Finally, allocation patterns are relatively stable when older females are better at acquiring food and are therefore in better condition. Our approach revealed an interaction between female state, age and survival costs, providing a novel explanation for observed variation in reproductive traits.     

Reproductive effort and costs of reproduction in female European ground squirrels

Reproductive effort, factors affecting reproductive output and costs of reproduction were studied in primiparous yearling compared to multiparous older female European ground squirrels (Spermophilus citellus). Yearling females weaned smaller litters than older ones. Litter size increased with posthibernation body mass at the expense of slightly lighter young for yearling but not for older mothers. In older females, on the other hand, emergence body mass influenced offspring mass, whereas litter size was affected by oestrus date. High reproductive effort entailed reproductive costs in terms of reduced subsequent fecundity but not subsequent survival for both yearling and older females. The production of large litters and long duration of lactation delayed subsequent oestrus, which, in turn, correlated negatively with litter size. During the second half of lactation, oestradiol levels were significantly elevated, indicating the initiation of follicular maturation processes. Oestradiol levels during that time correlated negatively with current, but positively with subsequent litter size. We therefore assume that inhibitory effects of lactation on gonadal development may mediate the negative relationship between reproductive effort and subsequent reproductive timing in adults. This effect is absent in yearlings because they are reproducing for the first time. Reproductive output in yearlings was influenced by interactions between structural growth and puberty. Received: 22 March 1999 / Accepted: 7 June 1999     

Effect of experimental ectoparasite control on bartonella infections in wild Richardson's ground squirrels

The purpose of this study was to investigate the role of ectoparasites in transmitting Bartonella infections in wild Richardson's ground squirrels (Spermophilus richardsonii). Richardson's ground squirrels were trapped, examined for fleas, and tested for Bartonella bacteremia once monthly, at six sites, from April to September 2004. After the initial trapping session in April, burrows at three sites were treated with deltamethrin insecticide. Richardson's ground squirrels trapped on treated sites were less likely to have fleas and had fewer fleas than squirrels on control sites in all months following treatment. We found no difference in the prevalence of Bartonella infections on control and treated sites in May, immediately following treatment; however, significantly fewer squirrels were infected with Bartonella on treated sites in June and July. We conclude that ectoparasites are a main route of transmission for Bartonella infections in Richardson's ground squirrels.     

Age‐specific responses to spring temperature in a migratory songbird: older females attempt more broods in warmer springs

Increasing global temperature has led to an interest in plasticity in the timing of annual events; however, little is known about the demographic consequences of changing phenology. Annual reproductive success varies significantly among individuals within a population, and some of that variation has to do with the number of broods attempted by reproducing adults. In birds, female age and the timing of reproduction are often predictors of multiple breeding. We hypothesize that double brooding rates may be affected by spring temperature and that the response may vary with female age. We used a long‐term reproductive data set for a migratory songbird, the prothonotary warbler (Protonotaria citrea) to assess which factors influence (a) an individual female's probability of double brooding and (b) the annual variation in population‐level double brooding rates. We found that older and earlier nesting birds are more likely to double brood, and that there is no evidence for senescence with regard to this trait such that the oldest females were most likely to double brood. Previous experience with double brooding (i.e., whether the female double brooded in the previous year) significantly increased the probability of doing so again. When assessing annual variation in the double brooding rate, we found an interaction between spring temperature and the proportion of older females in the population. Specifically, older females are more likely to double brood in years with warmer springs, but this relationship was not seen for younger females. Previous studies have shown that warmer temperatures lead to earlier and narrower peaks in resources and we hypothesize that these peaks are more available to older and earlier arriving females, enabling them to successfully raise more than one brood in a season. Understanding how different age classes respond to changing environmental conditions will be imperative to managing declining species.     

Is fatter fitter? Body storage and reproduction in ten populations of the freshwater amphipod Gammarus minus

Relationships between body storage (estimated as fat content and residuals of body mass regressed against body length) and offspring investment [brood mass, brood size (number of embryos per brood) and embryo mass] were examined within and among populations of the amphipod Gammarus minus in ten cold springs in central Pennsylvania, USA. Two major hypotheses and six corollary hypotheses were tested. Total reproductive investment (brood mass and brood size) was usually strongly positively correlated with maternal body length and body storage both within and among populations. These positive associations between reproductive and somatic investments are expected if individual variation in resource acquisition exceeds that of resource allocation. That is, individuals or populations that are able to acquire more resources should also be able to allocate more resources to both reproduction and somatic reserves than those acquiring fewer resources. This hypothesis is consistent with evidence showing that individual differences in body storage in G. minus and other amphipods are related to differences in resource acquisition. Positive associations between reproductive and somatic investments do not mean that energy costs of reproduction do not exist in G. minus. Evidence for reproductive energy costs included the lower body-fat contents of brooding versus nonbrooding females and the relatively low body mass per length of females who had just deposited eggs in their brood pouch. Unlike brood mass and brood size, individual embryo mass was usually unrelated to maternal body length and body storage. This pattern is largely consistent with optimal offspring investment theory, which predicts that offspring size should be insensitive to variation in parental resource status. However, in contrast to theory, embryo mass increased in winter when brooding females were significantly ”fatter”, presumably due to the availability of autumn-shed leaf food. This seasonal change in offspring size may be a maternally mediated effect of increased resource availability, though other explanations are possible. Overall, this study suggests that ”fatter” female amphipods are fitter than ”thinner” ones, though both the costs and benefits of increased body storage and brood size require investigation to substantiate this claim. This study also suggests that effects of individual variation in resource acquisition on life-history patterns deserve more theoretical and empirical attention by ecologists than they have received. It should be recognized that positive and/or nonsignificant correlations between life-history traits are just as interesting and important as are the negative correlations predicted by many theoretical models. Received: 20 January 1999 / Accepted: 26 September 1999     

Sex differences in senescence: the role of intra-sexual competition in early adulthood

Males and females frequently differ in their rates of ageing, but the origins of these differences are poorly understood. Sex differences in senescence have been hypothesized to arise, because investment in intra-sexual reproductive competition entails costs to somatic maintenance, leaving the sex that experiences stronger reproductive competition showing higher rates of senescence. However, evidence that sex differences in senescence are attributable to downstream effects of the intensity of intra-sexual reproductive competition experienced during the lifetime remains elusive. Here, we show using a 35 year study of wild European badgers (Meles meles), that (i) males show higher body mass senescence rates than females and (ii) this sex difference is largely attributable to sex-specific downstream effects of the intensity of intra-sexual competition experienced during early adulthood. Our findings provide rare support for the view that somatic maintenance costs arising from intra-sexual competition can cause both individual variation and sex differences in senescence.