Early-life behavior, survival, and maternal personality in a wild marsupial.

Individual behavior varies for many reasons, but how early in life are such differences apparent, and are they under selection? We investigated variation in early-life behavior in a wild eastern gray kangaroo (Macropus giganteus) population, and quantified associations of behavior with early survival. Behavior of young was measured while still in the pouch and as subadults, and survival to weaning was monitored. We found consistent variation between offspring of different mothers in levels of activity at the pouch stage, in flight initiation distance (FID) as subadults, and in subadult survival, indicating similarity between siblings. There was no evidence of covariance between the measures of behavior at the pouch young versus subadult stages, nor of covariance of the early-life behavioral traits with subadult survival. However, there was a strong covariance between FIDs of mothers and those of their offspring tested at different times. Further, of the total repeatability of subadult FID (51.5%), more than half could be attributed to differences between offspring of different mothers. Our results indicate that 1) behavioral variation is apparent at a very early stage of development (still in the pouch in the case of this marsupial); 2) between-mother differences can explain much of the repeatability (or "personality") of juvenile behavior; and 3) mothers and offspring exhibit similar behavioral responses to stimuli. However, 4) we found no evidence of selection via covariance between early-life or maternal behavioral traits and juvenile survival in this wild marsupial.

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.

Genetic rescue from protected areas is modulated by migration, hunting rate, and timing of harvest.

In terrestrial and marine ecosystems, migrants from protected areas may buffer the risk of harvest-induced evolutionary changes in exploited populations that face strong selective harvest pressures. Understanding the mechanisms favoring genetic rescue through migration could help ensure evolutionarily sustainable harvest outside protected areas and conserve genetic diversity inside those areas. We developed a stochastic individual-based metapopulation model to evaluate the potential for migration from protected areas to mitigate the evolutionary consequences of selective harvest. We parameterized the model with detailed data from individual monitoring of two populations of bighorn sheep subjected to trophy hunting. We tracked horn length through time in a large protected and a trophy-hunted populations connected through male breeding migrations. We quantified and compared declines in horn length and rescue potential under various combinations of migration rate, hunting rate in hunted areas and temporal overlap in timing of harvest and migrations, which affects the migrants' survival and chances to breed within exploited areas. Our simulations suggest that the effects of size-selective harvest on male horn length in hunted populations can be dampened or avoided if harvest pressure is low, migration rate is substantial, and migrants leaving protected areas have a low risk of being shot. Intense size-selective harvest impacts the phenotypic and genetic diversity in horn length, and population structure through changes in proportions of large-horned males, sex ratio and age structure. When hunting pressure is high and overlaps with male migrations, effects of selective removal also emerge in the protected population, so that instead of a genetic rescue of hunted populations, our model predicts undesirable effects inside protected areas. Our results stress the importance of a landscape approach to management, to promote genetic rescue from protected areas and limit ecological and evolutionary impacts of harvest on both harvested and protected populations.

Post-weaning survival in kangaroos is high and constant until senescence: Implications for population dynamics.

Large herbivores typically have consistently high prime-aged adult survival and lower, more variable, juvenile, and senescent survival. Many kangaroo populations undergo greater fluctuations in density compared with other large herbivores, but age- and sex-specific survival of kangaroos and their response to environmental variation remain poorly estimated. We used long-term capture-mark-recapture data on 920 individuals to investigate the survival component of eastern grey kangaroo (Macropus giganteus) population dynamics. Forage availability and population density were monitored quarterly and included as predictors of survival in Bayesian Cormack-Jolly-Seber models. Annual survival probabilities were estimated for five age classes: 0 years (juveniles), 1-2 years (subadults), 3-6 years (prime-aged adults), 7-9 years (presenescent adults), and ≥10 years (senescent adults). Survival of juveniles varied widely during our 12-year study, ranging from 0.07 to 0.90 for females and 0.05-0.92 for males. Subadult survival was 0.80-0.93 for females and 0.75-0.85 for males, while that of prime-aged adults was ≥0.94 for females and ≥0.83 for males, despite large fluctuations in forage and density. The survival of presenescent adults spanned 0.86-0.93 for females and 0.60-0.86 for males. Senescent survival was variable, at 0.49-0.90 for females and 0.49-0.80 for males. Male survival was significantly lower than female survival in prime-aged and presenescent adults, but not in other age classes. Although most of the models supported by Watanabe-Akaike Information Criterion selection included at least one environmental covariate, none of these covariates individually had a discernible effect on survival. Temporal variability in overall survival appeared mostly due to changes in the survival of juvenile and senescent kangaroos. Kangaroo survival patterns are similar to those of ungulates, suggesting a strong role of sex-age structure on population dynamics.

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.

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.

Genetic variance in fitness indicates rapid contemporary adaptive evolution in wild animals.

The rate of adaptive evolution, the contribution of selection to genetic changes that increase mean fitness, is determined by the additive genetic variance in individual relative fitness. To date, there are few robust estimates of this parameter for natural populations, and it is therefore unclear whether adaptive evolution can play a meaningful role in short-term population dynamics. We developed and applied quantitative genetic methods to long-term datasets from 19 wild bird and mammal populations and found that, while estimates vary between populations, additive genetic variance in relative fitness is often substantial and, on average, twice that of previous estimates. We show that these rates of contemporary adaptive evolution can affect population dynamics and hence that natural selection has the potential to partly mitigate effects of current environmental change.

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.

Breeding migrations by bighorn sheep males are driven by mating opportunities.

In some species where male mating success largely depends on intrasexual competition, males can adopt migratory or resident strategies to seek breeding opportunities. The resulting mixture of resident and migrant tactics within a population can have important ecological, genetic, and evolutionary consequences for metapopulations. Bighorn sheep Ovis canadensis males establish a linear dominance hierarchy that influences their mating tactics. Some males perform breeding migrations during the pre-rut and rut to seek mating opportunities, but little is known about these seasonal movements. We analyzed presence/absence data for 62 marked bighorn males during six mating seasons (20-32 males/year) in the Sheep River Provincial Park, Alberta, Canada, where hunting was not allowed. On average, about half of males left their natal population to rut elsewhere. The proportion of males leaving (yearly range 15%-69%) increased as the number of resident mature males increased and the populational sex ratio decreased, with fewer females during the pre-rut. Among those leaving the park, 24% did so in October, while the trophy sheep hunting season was open. Detailed monitoring of breeding migrations in protected populations could inform management strategies to limit evolutionary impacts of hunting, which can alter size-dependent mortality and create artificial pressures driving changes on heritable traits.

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.

Horn growth appears to decline under intense trophy hunting, but biases in hunt data challenge the interpretation of the evolutionary basis of trends.

A recent article in Evolutionary Applications by LaSharr et al. reports on trends in the size of horns of bighorn sheep (Ovis canadensis) throughout much of the species' range. The article concludes that there are "... stable or increasing trends in horn growth over nearly 3 decades in the majority of hunt areas throughout the western U.S. and Canada." However, the article equates nonsignificance of predominantly negative trends in the areas with the most selective harvest as evidence for the null hypothesis of no trends and also fails to consider well-known and serious biases in the use of data collected in size-regulated hunts. By applying meta-analysis to the estimates reported by LaSharr et al., we show that there has been a pervasive overall trend of declining horn sizes in Alberta, where the combination of horn size-based legality, combined with unrestricted hunter numbers are understood to generate the greatest selective pressures. Given the nature of the biases in the underlying data, the magnitudes of the trends resulting from our re-analysis of LaSharr et al.'s (Evolutionary Applications, 2019, 12, 1823) trend estimates are probably underestimated.

Determinants and long-term costs of early reproduction in males of a long-lived polygynous mammal.

In long-lived polygynous species, male reproductive success is often monopolized by a few mature dominant individuals. Young males are generally too small to be dominant and may employ alternative tactics; however, little is known about the determinants of reproductive success for young males. Understanding the causes and consequences of variability in early reproductive success may be crucial to assess the strength of sexual selection and possible long-term trade-offs among life-history traits. Selective pressures driven by fluctuating environmental conditions may depend on age class. We evaluated the determinants of reproduction in male bighorn sheep (Ovis canadensis) aged 2-4 years using 30 years of individual-level data. These young males cannot defend estrous ewes and use alternative mating tactics. We also investigated how the age of first detected reproduction was correlated to lifetime reproductive success and longevity. We found that reproductive success of males aged 3 years was positively correlated to body mass, to the proportion of males aged 2-4 years in the competitor pool, and to the number of females available per adult male. These results suggest that reproductive success depends on both competitive ability and population age-sex structure. None of these variables, however, had significant effects on the reproductive success of males aged 2 or 4 years. Known reproduction before the age of five increased lifetime reproductive success but decreased longevity, suggesting a long-term survival cost of early reproduction. Our analyses reveal that both individual-level phenotypic and population-level demographic variables influence reproductive success by young males and provide a rare assessment of fitness trade-offs in wild polygynous males.

Do Early-Life Conditions Drive Variation in Senescence of Female Bighorn Sheep?

The rate of senescence may vary among individuals of a species according to individual life histories and environmental conditions. According to the principle of allocation, changes in mortality driven by environmental conditions influence how organisms allocate resources among costly functions. In several vertebrates, environmental conditions during early life impose trade-offs in allocation between early reproduction and maintenance. The effects of conditions experienced during early life on senescence, however, remain poorly documented in wild populations. We examined how several early-life environmental conditions affected reproductive and survival senescence in wild bighorn sheep. We found long-term effects of high population density at birth, precipitations during the winter before birth, and temperature during the winter following birth that decreased survival after 7 years of age. High temperature during the first summer and autumn of life and high Pacific decadal oscillation decreased reproductive success at old ages. However, harsh early-life environment did not influence the rate of senescence in either survival or reproduction. Contrary to our expectation, we found no trade-off between reproductive allocation prior to senescence and senescence. Our results do show that early-life environmental conditions are important drivers of later survival and reproductive success and contribute to intra-specific variation in late-life fitness, but not aging patterns. These conditions should therefore be considered when studying the mechanisms of senescence and the determinants of variation in both survival and reproductive senescence at older ages.

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.

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.

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.

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.

Genetic decline, restoration and rescue of an isolated ungulate population.

Isolation of small populations is expected to reduce fitness through inbreeding and loss of genetic variation, impeding population growth and compromising population persistence. Species with long generation time are the least likely to be rescued by evolution alone. Management interventions that maintain or restore genetic variation to assure population viability are consequently of significant importance. We investigated, over 27 years, the genetic and demographic consequences of a demographic bottleneck followed by artificial supplementation in an isolated population of bighorn sheep (Ovis canadensis). Based on a long-term pedigree and individual monitoring, we documented the genetic decline, restoration and rescue of the population. Microsatellite analyses revealed that the demographic bottleneck reduced expected heterozygosity and allelic diversity by 6.2% and 11.3%, respectively, over two generations. Following supplementation, first-generation admixed lambs were 6.4% heavier at weaning and had 28.3% higher survival to 1 year compared to lambs of endemic ancestry. Expected heterozygosity and allelic diversity increased by 4.6% and 14.3% after two generations through new alleles contributed by translocated individuals. We found no evidence for outbreeding depression and did not see immediate evidence of swamping of local genes. Rapid intervention following the demographic bottleneck allowed the genetic restoration and rescue of this bighorn sheep population, likely preventing further losses at both the genetic and demographic levels. Our results provide further empirical evidence that translocation can be used to reduce inbreeding depression in nature and has the potential to mitigate the effect of human-driven environmental changes on wild populations.

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.