Lasting effects of conditions at birth on moose body mass

Whether cohort effects can be retrieved in adult phenotypes depends on the possibility for individuals to compensate for a good or bad start in life. This ability to compensate may itself depend on the environment and on individual sex. In large polygynous ungulates, male reproductive success relies more on body size than the reproductive success of females, which makes them more sensitive to a bad start in life. Based on current theories of life history evolution and sexual selection, we tested the following predictions in a moose population: 1) cohort effects and year effects occur in both male and female adult body mass, but due to 2) compensatory growth, cohort effects tend to fade away with the individual's age; and 3) males are more sensitive to cohort effects than females. In support of the first prediction, we found that density and climate during the year-of-birth and the year-of-harvest affected moose body mass in both sexes. However, the magnitude of the effects of environmental conditions at birth on adult body mass decreased with increasing age, but less so in males than in females. Thus, as expected based on our third prediction, environmental conditions early in life were more influential on adult body mass of males compared to females. Such a result supports the existence of sex-specific life history tactics, with males maximising growth rate and females rather trading-off growth for reproduction.     

Annual variation in maternal age and calving date generate cohort effects in moose (<Emphasis Type="Italic">Alces alces</Emphasis>) body mass

A general feature of the demography of large ungulates is that many demographic traits are dependent on female body mass at early ages. Thus, identifying the factors affecting body mass variation can give important mechanistic understanding of demographic processes. Here we relate individual variation in autumn and winter body mass of moose calves living at low density on an island in northern Norway to characteristics of their mother, and examine how these relationships are affected by annual variation in population density and climate. Body mass increased with increasing age of their mother, was lower for calves born late in the spring, decreased with litter size and was larger for males than for female calves. No residual effects of variation in density and climate were present after controlling for annual variation in mother age and calving date. The annual variation in adult female age structure and calving date explained a large part (71–75%) of the temporal variation in calf body mass. These results support the hypotheses that (a) body mass of moose calves are affected by qualities associated with mother age (e.g. body condition, calving date); and (b) populations living at low densities are partly buffered against temporal fluctuations in the environment.     

Life history patterns in female moose (Alces alces): the relationship between age,body size,fecundity and environmental conditions

I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.     

Catch-up growth in Japanese quail (Coturnix Japonica): relationships with food intake, metabolic rate and sex

The effects of early environmental conditions can profoundly affect individual development and adult phenotype. In birds, limiting resources can affect growth as nestlings, but also fitness and survival as adults. Following periods of food restriction, individuals may accelerate development, undergoing a period of rapid “catch-up” growth, in an attempt to reach the appropriate size at adulthood. Previous studies of altricial birds have shown that catch-up growth can have negative consequences in adulthood, although this has not been explored in species with different developmental strategies. Here, we investigated the effects of resource limitation and the subsequent period of catch-up growth, on the morphological and metabolic phenotype of adult Japanese quail (Coturnix japonica), a species with a precocial developmental strategy. Because males and females differ in adult body size, we also test whether food restriction had sex-specific effects. Birds that underwent food restriction early in development had muscles of similar size and functional maturity, but lower adult body mass than controls. There was no evidence of sex-specific sensitivity of food restriction on adult body mass; however, there was evidence for body size. Females fed ad lib were larger than males fed ad lib, while females subjected to food restriction were of similar size to males. Adults that had previously experienced food restriction did not have an elevated metabolic rate, suggesting that in contrast to altricial nestlings, there was no metabolic carry-over effect of catch-up growth into adulthood. While Japanese quail can undergo accelerated growth after re-feeding, timing of food restriction may be important to adult size, particularly in females. However, greater developmental flexibility compared to altricial birds may contribute to the lack of metabolic carryover effects at adulthood.     

Determinants of age at first reproduction and lifetime breeding success revealed by full paternity assignment in a male ungulate

Age at first reproduction is an important determinant of individual variation in reproductive success in ungulates, but few studies have examined its relationship with later fitness‐related traits in males. We used a long‐term individual based study of a harvested moose population to quantify the individual reproductive performance and survival of males, as well as to examine the determinants of age at first reproduction and consequences of age at first reproduction on lifetime breeding success. The probability that a male successfully reproduced at the age of two was negatively related to the mean age of adult males in the population, but the relationship weakened with increasing population size. Large antlers and large body mass relative to other males in the population increased the number of calves sired at their first successful mating season. In addition, those that successfully reproduced as two year‐olds were more likely to sire calves the next year, making them more productive at a given age compared to those that first reproduced at the age of three or older. We emphasize the importance for males to start reproducing as soon as possible in a harvested population to gain lifetime fitness benefits, as surviving the hunt is a major determinant of reproductive success in this population. We found no costs of early reproduction in males, hence leading to high individual heterogeneity in male reproductive performance. The apparent lack of reproductive costs could partly be explained by the age distribution in the population, individual variation in early‐life body mass and antler size, and differences in probabilities of being hunted of successful and unsuccessful males.     

Early growth in male and female fallow deer fawns

In this paper we present data from a long-term study on earlygrowth and related variables in fallow deer fawns living inlarge enclosures. Pre-winter body mass was constantly higherand more strongly correlated to subadult body mass in malesthan in females. To find out the mechanism for this higher pre-wintermass in males, we analyzed the variation in pre-winter massin relation to sex, year, mother's body mass, age and parityof mother, birth date, birth mass, growth rate, suckling behavior,and other behaviors. Birth mass was higher for male fawns, andgestation length, birth date, and weaning date did not differbetween the sexes. Consequently, both pre- and postnatal growthwere faster in males than in females. No behavioral differenceswere found between the sexes that could explain the differencein postnatal growth. Pre-winter mass was positively relatedto mother's body mass. Heavy mothers gave birth earlier andto larger offspring who grew at a higher rate, independent ofoffspring sex. However, male fawns born to primiparous mothershad relatively lower growth than male fawns born to multiparousmothers. This was not the case for female fawns. Suckling timeafter the first 2 weeks was positively related to mother's bodymass and growth of offspring. However, no measurements of sucklingbehavior differed between male and female fawns. Our results,except for the effect of parity on male and female growth, indicatethat selection has not acted on mothers to promote faster earlygrowth in males.     

Geographical variation in body weight and sexual size-dimorphism of Norwegian moose (Alces alces)

Geographical variation in male carcass weight, and sexual dimorphism in size was studied in 19 populations of Norwegian moose ( Alces alces (L.)).
Significant age-specific variation in male carcass weight was found for all the populations studied up to the age of 4 1/2 years, but in some populations maximum weight was not reached until at least 5 1/2 years. Increase in the mean weight of females after the age of 2 1/2 years was not significant. Only a weak relationship existed between mean yearling and adult bull weights in a population. However, within both the southern (< 62°N) and northern (> 62°N) parts of the country, yearling carcass weight was a good predictor of adult bull weight in a region.
Adult bull weight in a region was best predicted from the increment in mean carcass weight observed between 1 1/2 and 3 1/2 years of age. Within a region, variation in age-specific carcass weight between cohorts of bulls from different years was also well predicted from annual variation in growth increment.
Those patterns reveal a sexual difference in strategy of body growth. The adult weight of females is probably strongly determined by the weight gained by the time of onset of reproduction. The males have available a longer period for growth in body weight. They are therefore able to compensate for low weights early in life by increased gain of weight in later years, that provide good conditions for growth.
Geographical variation in the degree of sexual dimorphism in size correlated only poorly with adult male size. We suggest that the sexual size-dimorphism is a result of reproductive constraints of the female, i.e. in populations living in poor conditions and having small body size, the onset of reproduction prevents further gain in body weight.     

Age and sex-specific variation in detectability of moose (<Emphasis Type="Italic">Alces alces</Emphasis>) during the hunting season: implications for population monitoring

The sustainability of wild ungulate harvests can be greatly enhanced if monitoring data are available to permit an adaptive management approach. Utilising data provided by hunters is potentially the most cost-effective approach. In Scandinavia, observations recorded by moose (Alces alces) hunters provide a range of indices of population density, composition and reproductive performance. These are routinely used in practical management, but there are still many questions about their accuracy and precision. In this study, we availed of the fact that virtually all individual moose on the island of Vega in central Norway were marked during the period 1992–2005. Thus, we were able to compare the observation indices provided by hunters to the known size and composition of the population. The results indicate that the hunter observation indices provided accurate estimates of variation in moose cow recruitment and twinning rates. The estimates of sex ratio closely followed the pattern of annual variation but showed a consistent pattern of over-estimating the proportion of males. Thus, males seem to visually expose themselves more often for hunters than do females. The density index, measured as number of moose seen per hunter day did not to the same extent follow the variation in population density, possibly because moose were more naive in the first years after hunting was introduced or because of reduced detection conditions due to increasing scrub and tree encroachment during the study period. The results are discussed in light of their application to management and the evolutionary pressures in moose anti-predator behaviour.     

Size Matters: Individual Variation in Ectotherm Growth and Asymptotic Size

Body size, and, by extension, growth has impacts on physiology, survival, attainment of sexual maturity, fecundity, generation time, and population dynamics, especially in ectotherm animals that often exhibit extensive growth following attainment of sexual maturity. Frequently, growth is analyzed at the population level, providing useful population mean growth parameters but ignoring individual variation that is also of ecological and evolutionary significance. Our long-term study of Lake Erie Watersnakes, Nerodia sipedon insularum, provides data sufficient for a detailed analysis of population and individual growth. We describe population mean growth separately for males and females based on size of known age individuals (847 captures of 769 males, 748 captures of 684 females) and annual growth increments of individuals of unknown age (1,152 males, 730 females). We characterize individual variation in asymptotic size based on repeated measurements of 69 males and 71 females that were each captured in five to nine different years. The most striking result of our analyses is that asymptotic size varies dramatically among individuals, ranging from 631–820 mm snout-vent length in males and from 835–1125 mm in females. Because female fecundity increases with increasing body size, we explore the impact of individual variation in asymptotic size on lifetime reproductive success using a range of realistic estimates of annual survival. When all females commence reproduction at the same age, lifetime reproductive success is greatest for females with greater asymptotic size regardless of annual survival. But when reproduction is delayed in females with greater asymptotic size, lifetime reproductive success is greatest for females with lower asymptotic size when annual survival is low. Possible causes of individual variation in asymptotic size, including individual- and cohort-specific variation in size at birth and early growth, warrant further investigation.     

Population characteristics predict responses in moose body mass to temporal variation in the environment

1. A general problem in population ecology is to predict under which conditions stochastic variation in the environment has the stronger effect on ecological processes. By analysing temporal variation in a fitness-related trait, body mass, in 21 Norwegian moose Alces alces (L.) populations, we examined whether the influence of temporal variation in different environmental variables were related to different parameters that were assumed to reflect important characteristics of the fundamental niche space of the moose. 2. Body mass during autumn was positively related to early access to fresh vegetation in spring, and to variables reflecting slow phenological development (low June temperature, a long spring with a slow plant progression during spring). In contrast, variables related to food quantity and winter conditions had only a minor influence on temporal variation in body mass. 3. The magnitude of the effects of environmental variation on body mass was larger in populations with small mean body mass or living at higher densities than in populations with large-sized individuals or living at lower densities. 4. These results indicate that the strongest influence of environmental stochasticity on moose body mass occurs towards the borders of the fundamental niche space, and suggests that populations living under good environmental conditions are partly buffered against fluctuations in environmental conditions.     

Reproductive strategies and the influence of date of birth on growth and sexual development of an aseasonally-breeding ungulate: Reeves' muntjac (Muntiacus reevesi)

Unlike most species of deer, Reeves' muntjac does not have a seasonal reproductive cycle. Births are equally distributed throughout the year and, irrespective of season of birth, females start breeding when they have reached a minimum body weight of about 10 kg. A significantly smaller proportion of females is born in the autumn/winter than the spring/summer, but there is no effect of maternal age or condition on foetal sex ratios for first or subsequent pregnancies. Whilst aseasonal breeding increases female productivity, males are only able to hold territories encompassing the ranges of a number of does for a relatively short period of time. It is argued that for muntjac there is much less inter-sexual variation in lifetime reproductive success than for seasonally-breeding polygynous cervids. Hence, although muntjac are sexually dimorphic and polygynous, females do not invest preferentially in male offspring.
Young males can be fertile from 36 weeks of age, when their first antlers are still in velvet. Whilst season of birth has no effect on the rate of sexual development in females, it does for males, with autumn-born male fawns attaining sexual maturity earliest. This accelerated development occurs in the period following independence from the mother, and there is no evidence of a maternal cost in producing male fawns. Since the canine tusks are the main weapons for intra-sexual conflict, there could be clear gains in terms of reproductive success for males that attain sexual maturity, adult weight and adult tusk size quickly, even though their first antlers are much smaller than those of older bucks. The hypothesis is presented that, for muntjac, the majority of a male's lifetime reproductive success is achieved relatively early in life when the canine tusks are in pristine condition, and that bucks lose their territories, and associated access to oestrous does, once the canines are broken.     

根田鼠母体捕食应激效应与Trivers-Willard 模型的关系

本研究了根田鼠母体捕食应激对其子代出生、断乳和成体体重、窝性比及死亡率的作用,检验Trivers—Willard模型的2个前提条件、母体应激激素在母体投资中的作用,以及母体捕食应激效应与该模型的关系。将妊娠根田鼠母体暴露于其捕食艾鼬,母体应激子代的出生和断乳体重均显降低;到成体,雄性体重有此效应,而雌性体重则接近对照。出生窝性比无变化,但成体窝性比向雌性偏斜。不同年龄阶段的死亡率无显变化,但累计死亡率明显增大。处理雄性子代在断乳和成体时的皮质酮含量显增高,而雌性子代则无显变化,从而验证了Trivers-Willard模型的2个前提条件,提出应激母体激素参与母体对子代的投资观点,并认为,母体捕食应激使根田鼠子代成体窝性比向雌性偏斜的生理投资符合进化稳定对策。     

Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose

A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23–0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.     

Patterns of hunting mortality in Norwegian moose (Alces alces) populations

We used a simple life table approach to examine the age-specific patterns of harvest mortality in eight Norwegian moose populations during the last 15 years and tried to determine if the observed patterns were caused by hunter selectivity. The general opinion among local managers is that hunters prefer to shoot female moose not in company with calves to keep a high number of reproductive females in the population (and because of the emotional stress involved in leaving the calf/calves without a mother), and relatively large males because of the higher return with respect to meat and trophy. In support of the former view, we found the harvest mortality of adult females to be higher among pre-prime (1–3 years old) than prime-aged age classes (4–7 years old). This is probably because prime-aged females are more fecund and, therefore, more likely to be in company with one or two calves during the hunting season. As the season progressed, however, the selection pressure on barren females decreased, probably due to more productive females becoming ‘legal’ prey as their calf/calves were harvested. In males, we did not find any evidence of strong age-specific hunter selectivity, despite strong age-dependent variation in body mass and antler size. We suggest that this was due to the current strongly female-biased sex ratio in most Norwegian moose populations, which leaves the hunters with few opportunities to be selective within a relatively short and intensive hunting season. The management implications of these findings and to what extent the results are likely to affect the future evolution of life histories in Norwegian moose populations are discussed.Electronic Supplementary Material Supplementary material is available for this article at and accessible for authorised users.     

Maternal investment during pregnancy in wild meerkats

Maternal investment in offspring development is a major determinant of the survival and future reproductive success of both the mother and her young. Mothers might therefore be expected to adjust their investment according to ecological conditions in order to maximise their lifetime fitness. In cooperatively breeding species, where helpers assist breeders with offspring care, the size of the group may also influence maternal investment strategies because the costs of reproduction are shared between breeders and helpers. Here, we use longitudinal records of body mass and life history traits from a wild population of meerkats (Suricata suricatta) to explore the pattern of growth in pregnant females and investigate how the rate of growth varies with characteristics of the litter, environmental conditions, maternal traits and group size. Gestational growth was slight during the first half of pregnancy but was marked and linear from the midpoint of gestation until birth. The rate of gestational growth in the second half of pregnancy increased with litter size, maternal age and body mass, and was higher for litters conceived during the peak of the breeding season when it is hot and wet. Gestational growth rate was lower in larger groups, especially when litter size was small. These results suggest that there are ecological and physiological constraints on gestational growth in meerkats, and that females may also be able to strategically adjust their prenatal investment in offspring according to the likely fitness costs and benefits of a particular breeding attempt. Mothers in larger groups may benefit from reducing their investment because having more helpers might allow them to lower reproductive costs without decreasing breeding success.     

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.     

Life history of the moose Alces alces: relationship between growth and reproduction

We studied the relationship between increase in body weight and reproductive performance in different populations of Norwegian moose to evaluate costs associated with early onset of reproduction, viz. whether early onset of reproduction was correlated with low adult body weight or reduced adult fecundity. The mean carcass weight of non-ovulating yearlings was significantly lower than for ovulating yearlings. However, those 2.5 yr old females that conceived as yearlings were lighter than non-reproducing females of the same age. Thus, to begin to reproduce as a yearling was assumed to be expensive because it reduces the possibilities for further growth. The cost associated with reproduction was further illustrated by the fact that the difference in mean carcass weight from age 2.5 to 3.5 yr of females that produced calves in both years, was less for the females from regions with lowest mean yearling weights, i.e. regions with probably the lowest resource availability. In populations with high mean yearling carcass weights, the proportion of cows with calf and the number of calves per pregnant female in the early reproductive phase (2.5 or 3.5 yr old) were higher than in populations in which the mean yearling weights were low. There was a negative correlation between growth rate in the population after 1.5 yr of age and the mean yearling carcass weight. Thus, low yearling weight was associated with a prolonged period of growth and thereby a reduced reproductive output during the first year of the female's life. For old females (≥ 9.5 yr old) the number of calves produced per pregnant female was highest in populations where yearling carcass weights were highest. Furthermore, mean yearling weight and the mean adult female weight were positively correlated in those regions. This relationship suggests that within this species early onset of reproduction is not related to retarded reproduction or lower weight later in life. We suggest that the moose has been selected for an early onset of reproduction.     

Seasonality in Reproduction of <Emphasis Type="Italic">Lepilemur edwardsi</Emphasis>

Sportive lemurs are threatened species in the IUCN Red Data Book. However, quantitative information on their reproduction, urgently required for conservation, has been lacking. We collected first data on reproductive activity of Milne-Edwards’ sportive lemurs in a population inhabiting the dry deciduous forest of the Ankarafantsika National Park in northwestern Madagascar during 1998, 2001, and 2003. The species showed a seasonal reproduction. The main mating season extended from May to June, as indicated by the presence of males with high testes volumes and estrus females. In the mating and early postmating season and in the postparturition season, sexes did not differ in body mass. Females had a significantly higher body mass than males in August and November, indicating pregnancy, which together with the presence of small infants in October and November implies that gravidity in females lasted for about 4–5 mo. All litters consisted of singletons. Individuals with body mass at the lower limit of the population either did not develop measurable testes volumes (males) or were not in estrus (females). They were probably juveniles from the previous birth season that achieved sexual maturity not before their second year after birth. The first data on reproduction suggest a low reproduction rate for Lepilemur edwardsi and a request for a higher conservation status than previously attributed and the need for further management strategies.     

Effects of inbreeding on fitness‐related traits in a small isolated moose population

Inbreeding can affect fitness‐related traits at different life history stages and may interact with environmental variation to induce even larger effects. We used genetic parentage assignment based on 22 microsatellite loci to determine a 25 year long pedigree for a newly established island population of moose with 20–40 reproducing individuals annually. We used the pedigree to calculate individual inbreeding coefficients and examined for effects of individual inbreeding (f) and heterozygosity on fitness‐related traits. We found negative effects of f on birth date, calf body mass and twinning rate. The relationship between f and calf body mass and twinning rate were found to be separate but weaker after accounting for birth date. We found no support for an inbreeding effect on the age‐specific lifetime reproductive success of females. The influence of f on birth date was related to climatic conditions during the spring prior to birth, indicating that calves with a low f were born earlier after a cold spring than calves with high f. In years with a warm spring, calf f did not affect birth date. The results suggest that severe inbreeding in moose has both indirect effects on fitness through delayed birth and lower juvenile body mass, as well as separate direct effects, as there still was a significant relationship between f and twinning rate after accounting for birth date and body mass as calf. Consequently, severe inbreeding as found in the study population may have consequences for population growth and extinction risk.     

Determinants and consequences of age of primiparity in bighorn ewes

Because variation in age of first reproduction can have major effects on individual fitness and population dynamics, it is important to understand what maintains that variability. Although early primiparity is assumed to be costly, it is sometimes associated with high lifetime reproductive success. We used a long‐term study on bighorn sheep Ovis canadensis to determine what variables affect age at first reproduction, investigate the impact of primiparity on body resources and quantify the reproductive performance of primiparous ewes. We then examined the consequences of delayed primiparity on adult body mass, longevity and lifetime reproductive success. Environmental conditions during early development, body mass as a yearling, genotype and maternal effects affected age of primiparity. Primiparous ewes lost more mass in winter and gained less mass in summer than multiparous ewes. Small yearling ewes that postponed reproduction attained similar adult mass than heavy yearling ewes who reproduced at a younger age. Early primiparity did not reduce longevity and was positively associated with lifetime reproductive success. Starting to reproduce as soon as possible appears to maximize fitness of females. When early life conditions are unfavorable, however, delayed primiparity allows greater body growth and likely maximizes survival. The combination of a conservative reproductive strategy and maternal effects on age of primiparity may partly delay population recovery following density‐dependent declines.