Birth date determines early calf survival in migratory caribou.

The decline of most caribou (Rangifer tarandus) populations underlines the need to understand the determinants of key demographic parameters. In migratory caribou, we have limited information on rates and drivers of pre-weaning mortality. We fitted 60 pregnant females of the Rivière-aux-Feuilles caribou herd with GPS camera collars to track the survival of calves from birth to weaning in 2016-2018. Over the three years, calf survival rate before weaning, i.e. to 01-Sep, approximately three months of age, was 0.63 (CI 0.50-0.77). Summer mortality risk was mainly influenced by calf birth date, with calves born earlier in the calving season having a lower mortality risk than those born later. Mortality also increased when calves experienced low or high temperature during calving. This study provides the first estimates of pre-weaning survival of migratory caribou calves in this herd, illustrating the value of new technologies to collect data otherwise difficult to obtain in widely distributed migratory populations. This approach can easily be extended to other large herbivores and predators. Our study brings new insights on how climate change may affect summer juvenile survival given the increased temperatures and faster changes in plant phenology expected in the future.

Sons accelerate maternal aging in a wild mammal.

Aging, or senescence, is a progressive deterioration of physiological function with age. It leads to age-related declines in reproduction (reproductive senescence) and survival (actuarial senescence) in most organisms. However, senescence patterns can be highly variable across species, populations, and individuals, and the reasons for such variations remain poorly understood. Evolutionary theories predict that increases in reproductive effort in early life should be associated with accelerated senescence, but empirical tests have yielded mixed results. Although in sexually size-dimorphic species offspring of the larger sex (typically males) commonly require more parental resources, these sex differences are not currently incorporated into evolutionary theories of aging. Here, we show that female reproductive senescence varies with both the number and sex ratio of offspring weaned during early life, using data from a long-term study of bighorn sheep. For a given number of offspring, females that weaned more sons than daughters when aged between 2 and 7 y experienced faster senescence in offspring survival in old age. By contrast, analyses of actuarial senescence showed no cost of early-life reproduction. Our results unite two important topics in evolutionary biology: life history and sex allocation. Offspring sex ratio may help explain among-individual variation in senescence rates in other species, including humans.

Fluctuating optimum and temporally variable selection on breeding date in birds and mammals.

Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection.

Kangaroo Fathers Modulate Maternal Control of Offspring Sex but Not Postpartum Maternal Expenditure.

AbstractWhen sons and daughters have different fitness costs and benefits, selection may favor deviations from an even offspring sex ratio. Most theories on sex ratio manipulation focus on maternal strategies and sex-biased maternal expenditure. Recent studies report paternal influences on both offspring sex ratio and postpartum sex-biased maternal expenditure. We used long-term data on marked kangaroos to investigate whether and how paternal mass and skeletal size, both determinants of male reproductive success, influenced (a) offspring sex in interaction with maternal mass and (b) postpartum sex-biased maternal expenditure. When mothers were light, the probability of having a son increased with paternal mass. Heavy mothers showed the opposite trend. A similar result emerged when considering paternal size instead of mass. Postpartum maternal sex-specific expenditure was independent of paternal mass or size. Studies of offspring sex manipulation or maternal expenditure would benefit from an explicit consideration of paternal traits, as paternal and maternal effects can modulate each other.

Ewe are what ewe wear: bigger horns, better ewes and the potential consequence of trophy hunting on female fitness in bighorn sheep.

In polygynous species, secondary sexual traits such as weapons or elaborate ornaments have evolved through intrasexual competition for mates. In some species, these traits are present in both sexes but are underdeveloped in the sex facing lower intrasexual competition for mates. It is often assumed that these underdeveloped sexually selected traits are a vestige of strong sexual selection on the other sex. Here, we challenge this assumption and investigate whether the expression of secondary sexual traits is associated with fitness in female bighorn sheep. Analyses of 45 years of data revealed that female horn length at 2 years, while accounting for mass and environmental variables, is associated with younger age at primiparity, younger age of first offspring weaned, greater reproductive lifespan and higher lifetime reproductive success. There was no association between horn length and fecundity. These findings highlight a potential conservation issue. In this population, trophy hunting selects against males with fast-growing horns. Intersexual genetic correlations imply that intense selective hunting of large-horned males before they can reproduce can decrease female horn size. Therefore, intense trophy hunting of males based on horn size could reduce female reproductive performance through the associations identified here, and ultimately reduce population growth and viability.

Testing the match-mismatch hypothesis in bighorn sheep in the context of climate change.

In species with long gestation, females commit to reproduction several months before parturition. If cues driving conception date are uncoupled from spring conditions, parturition could be mistimed. Mismatch may increase with global change if the rate of temporal changes in autumn cues differs from the rate of change in spring conditions. Using 17 years of data on climate and vegetation phenology, we show that autumn temperature and precipitation, but not vegetation phenology, explain parturition date in bighorn sheep. Although autumn cues drive the timing of conception, they do not predict conditions at parturition in spring. We calculated the mismatch between individual parturition date and spring green-up, assessed whether mismatch increased over time and investigated the consequences of mismatch on lamb neonatal survival, weaning mass and overwinter survival. Mismatch fluctuated over time but showed no temporal trend. Temporal changes in green-up date did not lead to major fitness consequence of mismatch. Detailed data on individually marked animals revealed no effect of mismatch on neonatal or overwinter survival, but lamb weaning mass was negatively affected by mismatch. Capital breeders might be less sensitive to mismatch than income breeders because they are less dependent on daily food acquisition. Herbivores in seasonal environments may access sufficient forage to sustain lactation before or after the spring 'peak' green-up, and partly mitigate the consequences of a mismatch. Thus, the effect of phenological mismatch on fitness may be affected by species life history, highlighting the complexity in quantifying trophic mismatches in the context of climate change.

Determinants of spring molt in bighorn sheep: life-history, plasticity and phenology.

Anthropogenic climate change and habitat alterations increase the importance of understanding the causes and consequences of variation in phenological traits. Although the timing of phenological events may vary in response to both direct and mediated effects, methods to measure and distinguish direct and mediated effects have seldom been used. We used a Bayesian structural equation model (SEM) to evaluate potential direct and mediated effects of intrinsic individual and environmental factors on the timing and progression of spring molt in bighorn sheep. The SEM showed that molt phenology varied across years, was earlier in prime-aged and in heavier individuals, slower in males, and later in lactating ewes, especially if they were light. These results highlight how individual variation in intrinsic traits and life-history leads to substantial variation in a phenological trait. Indirect effects in the SEM predicted a delay in sheep molt phenology at high population density mediated through negative density effects on body mass and lactation probability. Cooler temperatures in late spring were also predicted to delay molt phenology via a negative effect on body mass. Finally, lactation reduced ewe mass which was predicted to delay molt phenology. This mediated effect thus increased the total delay (sum of direct and mediated effects) in molt phenology experience by lactating ewes. Our results underline the importance of estimating direct and indirect effects when modeling phenological traits. Because indirect effects could substantially affect estimates of total plasticity, they should be critically important to accurately predict phenological mismatches and demographic consequences of environmental change.

Correction to: Forage availability and maternal characteristics affect costs of reproduction in a large marsupial.

Unfortunately, the online publication contained an error in the "Data availability statement" and it is corrected by this erratum.

Forage availability and maternal characteristics affect costs of reproduction in a large marsupial.

Life history theory predicts trade-offs in allocation between survival, maintenance, growth, and reproduction, especially when resources are scarce. Individual variation in resource acquisition can affect trade-offs, but is often unaccounted for. We quantified the fitness costs of reproduction, accounting for environmental conditions, maternal characteristics and individual variation. We analyzed 10 years of data from marked kangaroos to evaluate how reproductive allocation affected annual mass change and skeletal growth, subsequent fecundity and weaning success, and survival, accounting for maternal mass or size and forage availability. Through repeated measurements of 76-91 females, we investigated how trade-offs varied within and between individuals, assessing whether individual variation could mask population-level trade-offs. In poor environments, females that weaned an offspring lost mass. Females that nursed an offspring for > 7 months had reduced skeletal growth. Females that did not gain mass over the previous 12 months rarely reproduced, especially if they had nursed an offspring for > 7 months the previous year. Reproductive allocation had no effect on weaning success, which was very low, and did not affect maternal survival, suggesting a conservative strategy. Disentangling within- and between-individual responses revealed trade-offs within individuals, but because individuals did not vary in their responses to earlier effort, these trade-offs did not drive population trends. The interacting effects of environmental conditions, maternal characteristics and individual variation on allocation trade-offs demonstrate the importance of long-term monitoring for understanding life history variations in changing environments.

Milk composition in a wild mammal: a physiological signature of phenological changes.

Understanding how spring phenology influences early life can provide important insights into drivers of future development and survival. We combined unique, long-term data from a bighorn sheep population and satellite-derived phenology indices to quantify the relative importance of maternal and environmental influences on milk composition and lamb overwinter survival. Based on 216 milk samples from 34 females monitored over 6 years, we found that longer snow-free and vegetation growing seasons increased milk fatty acid, iron and lactose concentrations. Structural equation modelling revealed no causality between milk energy content, lamb weaning mass and lamb overwinter survival. Our results suggest that spring conditions can affect milk energy content, but we did not detect any effect on lamb overwinter survival either directly or indirectly through lamb weaning mass. The effect of green-up date on milk composition and energy content suggests that herbivores living in seasonal environments, such as the bighorn sheep, might rely on a strategy intermediate between 'capital' and 'income' breeding when energy demands are high.

Maternal longevity and offspring sex in wild ungulates.

In species with sexual size dimorphism, the offspring of the larger sex usually have greater energy requirements and may lead to greater fitness costs for parents. The effects of offspring sex on maternal longevity, however, have only been tested in humans. Human studies produced mixed results and considerable debate mainly owing to the difficulty of distinguishing the effects of sexual dimorphism from sociocultural factors. To advance this debate, we examined how the relative number of sons influenced maternal longevity in four species of free-living ungulates (Soay sheep Ovis aries; bighorn sheep, Ovis canadensis; red deer, Cervus elaphus; mountain goat, Oreamnos americanus), with high male-biased sexual size dimorphism but without complicating sociocultural variables. We found no evidence for a higher cumulative cost of sons than of daughters on maternal longevity. For a given number of offspring, most females with many sons in all four populations lived longer than females with few sons. The higher cost of sons over daughters on maternal lifespan reported by some human studies may be the exception rather than the rule in long-lived iteroparous species.

Trophy hunting mediates sex-specific associations between early-life environmental conditions and adult mortality in bighorn sheep.

Environmental conditions during early development, from conception to sexual maturity, can have lasting consequences on fitness components. Although adult life span often accounts for much of the variation in fitness in long-lived animals, we know little about how early environment affects adult life span in the wild, and even less about whether these effects differ between the sexes. Using data collected over 45 years from wild bighorn sheep (Ovis canadensis), we investigated the effects of early environment on adult mortality in both sexes, distinguishing between natural and anthropogenic sources of mortality. We used the average body mass of yearlings (at about 15 months of age) as a yearly index of environmental quality. We first examined sex differences in natural mortality responses to early environment by censoring harvested males in the year they were shot. We then investigated sex differences in the effects of early environment on overall mortality (natural and hunting mortality combined). Finally, we used path analysis to separate the direct influence of early environment from indirect influences, mediated by age at first reproduction, adult mass and horn length. As early environmental conditions improved, natural adult mortality decreased in both sexes, although for males the effect was not statistically supported. Sex differences in the effects of early environment on adult mortality were detected only when natural and hunting mortality were pooled. Males that experienced favourable early environment had longer horns as adults and died earlier because of trophy hunting, which does not mimic natural mortality. Females that experienced favourable early environment started to reproduce earlier and early primiparity was associated with reduced mortality, suggesting a silver-spoon effect. Our results show that early conditions affect males and females differently because of trophy hunting. These findings highlight the importance of considering natural and anthropogenic environmental factors across different life stages to understand sex differences in mortality.

Long-term studies of bighorn sheep and mountain goats reveal fitness costs of reproduction.

Fitness costs of reproduction are expected when resources are limited. Costs drive the evolution of life-history strategies and can affect population dynamics if females change their allocation of resources to reproduction. We studied fitness costs of reproduction in mountain ungulates in Alberta, Canada. We monitored two populations of bighorn sheep (Ovis canadensis) for 44 and 30 years, and one of mountain goats (Oreamnos americanus) for 30 years. Both species are highly iteroparous. Heterogeneity in individual reproductive potential makes fitness costs of reproduction difficult to detect and quantify without manipulations. In capital breeders, individual differences can be partly accounted for by considering body mass and other correlates of reproductive potential. Long-term monitoring can reveal costs that only manifest under stressful conditions such as disease or resource scarcity. Despite individual differences in reproductive potential, we detected fitness costs of reproduction in females. Costs, in terms of mass gain and survival, are almost entirely born by subsequent offspring, as mothers prioritize their own maintenance and survival. Costs are greater for primiparous females, decrease with increasing body mass and increase as resource availability declines, and sons are costlier than daughters. Costs may increase for senescent females that appear to reduce allocation to reproduction. In bighorn sheep, costs mostly involve reduced mass gain and lower survival of subsequent offspring. In mountain goats, costs include reductions in mass gain, subsequent fecundity and juvenile survival. In males, fitness costs derive mostly from attempts to reproduce rather than from siring success and likely depend upon individual competitiveness. In the absence of selective harvests, dominant males may enjoy high fitness and possibly lower costs compared to subordinates. The conservative reproductive tactic of mountain ungulate females likely explains why density dependence mostly involves later primiparity and lower recruitment, but rarely affects adult survival. Future research will seek to better account for heterogeneity in reproductive potential, assess cumulative reproductive costs and investigate the potential effects of fathers on maternal allocation tactics.

Les coûts de la reproduction émergent lorsque les ressources sont limitées et influencent l'évolution des stratégies d'histoire de vie. Si les femelles modifient l'allocation des ressources dans la reproduction pour mitiger ces coûts, ils peuvent aussi affecter la dynamique de population. Nous avons étudié les coûts de la reproduction chez des ongulés en Alberta, Canada. Nous avons suivi deux populations de mouflons d'Amérique (Ovis canadensis) pour 44 et 30 ans, et une de chèvre de montagne (Oreamnos americanus) pour 30 ans. Ces deux espèces sont itéropares. L'hétérogénéité dans le potentiel reproducteur des individus peut masquer les coûts en fitness liés à la reproduction. Ils sont donc plus souvent détectables à l'aide de manipulation expérimentale. Chez les espèces avec reproduction « sur capital ¼, il est possible de contrôler pour les différences individuelles en incluant des variables liées au potentiel reproducteur telles que la masse corporelle. Les suivis à long terme permettent de détecter des coûts qui se manifestent seulement lors de conditions environnementales défavorables. Malgré les différences en potentiel reproducteur, nous avons détecté des coûts chez les femelles. Ces coûts, tels qu'une réduction en croissance ou en survie, sont presque toujours subis par les jeunes nés lors d'épisodes de reproduction subséquents. En effet, les mères priorisent leur propre maintien et leur survie. Les coûts sont supérieurs pour les femelles primipares et dans les conditions défavorables. Ils sont plus faibles pour les femelles plus lourdes et il semble que les fils soient plus coûteux que les filles. Les coûts augmentent aussi chez les femelles sénescentes qui semblent réduire leur allocation dans la reproduction. Chez les mouflons, les coûts impliquent une réduction du gain en masse et une survie plus faible des jeunes nés lors de reproductions subséquentes. Chez les chèvres de montagne, ces coûts sont accompagnés d'une réduction de la fécondité future. Chez les mâles, les coûts sont surtout influencés par l'investissement en temps et en énergie dans le rut plutôt que par le succès d'accouplement. Ces coûts devraient donc surtout dépendre de l'habileté compétitive individuelle. En absence de chasse sélective, les mâles dominants devraient avoir un fort succès reproducteur avec des coûts potentiellement plus faibles comparativement aux subordonnées. La tactique conservative adoptée par les ongulés de montagne femelles peut expliquer pourquoi les effets densité-dépendant mènent à un retard dans l'âge à la primiparité et à un recrutement plus faible, mais affectent rarement la survie adulte. Nos recherches futures tenteront de mieux quantifier l'hétérogénéité individuelle, évalueront les coûts cumulatifs à la reproduction et investigueront l'effet potentiel des pères sur les tactiques d'allocation maternelles.

Drivers and demographic consequences of seasonal mass changes in an alpine ungulate.

We know little about the determinants and demographic consequences of the marked seasonal mass changes exhibited by many northern and alpine mammals. We analysed 43 years of data on individual winter mass loss (the difference between mass in early June and mass in mid-September the previous year) and summer mass gain (the difference between mass in mid-September and in early June of the same year) in adult bighorn sheep (Ovis canadensis). We calculated relative seasonal mass change as a proportion of individual body mass at the start of each season. We first examined the effects of weather and population density on relative changes in body mass. We then assessed the consequences of relative seasonal mass changes on reproduction. Mean April-May temperature was the main driver of relative seasonal mass changes: warm springs reduced both relative winter mass loss and summer mass gain of both sexes, likely partially due to a trade-off between growth rate of plants and duration of access to high-quality forage. Because these effects cancelled each other, spring temperature did not influence mass in mid-September. Mothers that lost relatively more mass during the winter had lambs that gained less mass during summer, likely because these females allocated fewer resources to lactation. Winter survival of lambs increased with their summer mass gain. In males, relative mass loss during winter, which includes the rut, did not influence the probability of siring at least one lamb, possibly indicating that greater mating effort did not necessarily translate into greater reproductive success. Our findings improve our understanding of how weather influences recruitment and underline the importance of cryptic mechanisms behind the effects of climate change on demographic traits.

Tall young females get ahead: size-specific fecundity in wild kangaroos suggests a steep trade-off with growth.

When resources are limited, organisms face allocation conflicts. Indeterminate growth creates a persistent conflict with reproduction, as growth may enhance future reproduction, but diverts resources from current reproduction. Little is known about allocation trade-offs in mammals with indeterminate growth. We studied growth and reproduction in adult female eastern grey kangaroos (Macropus giganteus), an iteroparous mammal with indeterminate growth. Allocation trajectories varied with age and size: for 4-year-old females, fecundity increased from 30 to 82% from shortest to average-sized individuals. Older females had high fecundity regardless of size. The smallest females grew 30% more annually than average-sized females, but females that reached average size at an older age had lower growth rates. Environmental conditions affected allocation to size and reproduction. Rainy springs increased fecundity from 61 to 84% for females that had previously reproduced, but rainy winters reduced leg growth. Females in better relative condition grew 40% more than average, whereas most young of females below average relative condition failed to survive to 10 months of age. These results highlight an age-specific trade-off between growth and reproduction. Tall young females benefit from a smaller trade-off between somatic growth and early fecundity than shorter females of the same age, but older females appear to favor reproduction over growth regardless of size. Our study highlights how individual heterogeneity determines trade-offs between life-history components. We speculate that cohort effects affect age-specific reproductive success in this long-lived mammal.

Long-term fitness consequences of early environment in a long-lived ungulate.

Cohort effects can be a major source of heterogeneity and play an important role in population dynamics. Silver-spoon effects, when environmental quality at birth improves future performance regardless of the adult environment, can induce strong lagged responses on population growth. Alternatively, the external predictive adaptive response (PAR) hypothesis predicts that organisms will adjust their developmental trajectory and physiology during early life in anticipation of expected adult conditions but has rarely been assessed in wild species. We used over 40 years of detailed individual monitoring of bighorn ewes (Ovis canadensis) to quantify long-term cohort effects on survival and reproduction. We then tested both the silver-spoon and the PAR hypotheses. Cohort effects involved a strong interaction between birth and current environments: reproduction and survival were lowest for ewes that were born and lived at high population densities. This interaction, however, does not support the PAR hypothesis because individuals with matching high-density birth and adult environments had reduced fitness. Instead, individuals born at high density had overall lower lifetime fitness suggesting a silver-spoon effect. Early-life conditions can induce long-term changes in fitness components, and their effects on cohort fitness vary according to adult environment.

Fluctuating effects of genetic and plastic changes in body mass on population dynamics in a large herbivore.

Recent studies suggest that evolutionary changes can occur on a contemporary time scale. Hence, evolution can influence ecology and vice-versa. To understand the importance of eco-evolutionary dynamics in population dynamics, we must quantify the relative contribution of ecological and evolutionary changes to population growth and other ecological processes. To date, however, most eco-evolutionary dynamics studies have not partitioned the relative contribution of plastic and evolutionary changes in traits on population, community, and ecosystem processes. Here, we quantify the effects of heritable and non-heritable changes in body mass distribution on survival, recruitment, and population growth in wild bighorn sheep (Ovis canadensis) and compare their importance to the effects of changes in age structure, population density, and weather. We applied a combination of a pedigree-based quantitative genetics model, statistical analyses of demography, and a new statistical decomposition technique, the Geber method, to a long-term data set of bighorn sheep on Ram Mountain (Canada), monitored individually from 1975 to 2012. We show three main results: (1) The relative importance of heritable change in mass, non-heritable change in mass, age structure, density, and climate on population growth rate changed substantially over time. (2) An increase in body mass was accompanied by an increase in population growth through higher survival and recruitment rate. (3) Over the entire study period, changes in the body mass distribution of ewes, mostly through non-heritable changes, affected population growth to a similar extent as changes in age structure or in density. The importance of evolutionary changes was small compared to that of other drivers of changes in population growth but increased with time as evolutionary changes accumulated. Evolutionary changes became increasingly important for population growth as the length of the study period considered increased. Our results highlight the complex ways in which ecological and evolutionary changes can affect population dynamics and illustrate the large potential effect of trait changes on population processes.

Changes in horn size of Stone's sheep over four decades correlate with trophy hunting pressure.

Selective harvest may lead to rapid evolutionary change. For large herbivores, trophy hunting removes males with large horns. That artificial selection, operating in opposition to sexual selection, can lead to undesirable consequences for management and conservation. There have been no comparisons of long-term changes in trophy size under contrasting harvest pressures. We analyzed horn measurements of Stone's rams (Ovis dalli stonei) harvested over 37 years in two large regions of British Columbia, Canada, with marked differences in hunting pressure to identify when selective hunting may cause a long-term decrease in horn growth. Under strong selective harvest, horn growth early in life and the number of males harvested declined by 12% and 45%, respectively, over the study period. Horn shape also changed over time: horn length became shorter for a given base circumference, likely because horn base is not a direct target of hunter selection. In contrast, under relatively lower hunting pressure, there were no detectable temporal trends in early horn growth, number of males harvested, or horn length relative to base circumference. Trophy hunting is an important recreational activity and can generate substantial revenues for conservation. By providing a reproductive advantage to males with smaller horns and reducing the availability of desirable trophies, however, excessive harvest may have the undesirable long-term consequences of reducing both the harvest and the horn size of rams. These consequences can be avoided by limiting offtake.

Maternal condition and previous reproduction interact to affect offspring sex in a wild mammal.

Trivers and Willard proposed that offspring sex ratio should vary with maternal condition when condition, meant as maternal capacity to care, has different fitness consequences for sons and daughters. In polygynous and dimorphic species, mothers in good condition should preferentially produce sons, whereas mothers in poor condition should produce more daughters. Despite its logical appeal, support for this hypothesis has been inconsistent. Sex-ratio variation may be influenced by additional factors, such as environmental conditions and previous reproduction, which are often ignored in empirical studies. We analysed 39 years of data on bighorn sheep (Ovis canadensis) that fit all the assumptions of the Trivers-Willard hypothesis. Production of sons increased with maternal condition only for mothers that weaned a son the previous year. This relationship likely reflects a mother's ability to bear the higher reproductive costs of sons. The interaction between maternal condition and previous weaning success on the probability of producing a son was independent of the positive effect of paternal reproductive success. Maternal and paternal effects accounted for similar proportions of the variance in offspring sex. Maternal reproductive history should be considered in addition to current condition in studies of sex allocation.

Individual heterogeneity and offspring sex affect the growth-reproduction trade-off in a mammal with indeterminate growth.

Reproduction can lead to a trade-off with growth, particularly when individuals reproduce before completing body growth. Kangaroos have indeterminate growth and may always face this trade-off. We combined an experimental manipulation of reproductive effort and multi-year monitoring of a large sample size of marked individuals in two populations of eastern grey kangaroos to test the predictions (1) that reproduction decreases skeletal growth and mass gain and (2) that mass loss leads to reproductive failure. We also tested if sex-allocation strategies influenced these trade-offs. Experimental reproductive suppression revealed negative effects of reproduction on mass gain and leg growth from 1 year to the next. Unmanipulated females, however, showed a positive correlation between number of days lactating and leg growth over periods of 2 years and longer, suggesting that over the long term, reproductive costs were masked by individual heterogeneity in resource acquisition. Mass gain was necessary for reproductive success the subsequent year. Although mothers of daughters generally lost more mass than females nursing sons, mothers in poor condition experienced greater mass gain and arm growth if they had daughters than if they had sons. The strong links between individual mass changes and reproduction suggest that reproductive tactics are strongly resource-dependent.