Variation In Lifetime Reproductive Success Research Articles

The Ne/N ratio in applied conservation.

Recent developments within the IUCN and the Convention on Biological Diversity have affirmed the increasingly key role that effective population size (N e) and the effective size: census size ratio (N e/N) play in applied conservation and management of global biodiversity. This paper reviews and synthesizes information regarding the definition of N e and demographic and genetic methods for estimating effective size, census size, and their ratio. Emphasis is on single-generation estimates of contemporary N e/N, which are the most informative for practical applications. It is crucial to clearly define which individuals are included in the census size (N). Defining N as the number of adults alive at a given time facilitates comparisons across species. For a wide range of applications and experimental designs, inbreeding N e is simpler to calculate and interpret than variance N e. Effects of skewed sex ratio are generally modest, so most reductions to N e/N arise from overdispersed (greater-than-Poisson) variance in offspring number (). Even when fecundity changes with age, overdispersed within-age variance generally contributes most to overall , and both random and deterministic (mediated by selection) factors can be important. Most species are age-structured, so it is important to distinguish between effective size per generation (N e) and the effective number of breeders in one season or year (N b). Both N e and N b are important for applied conservation and management. For iteroparous species, a key metric is variance in lifetime reproductive success (), which can be affected by a variety of additional factors, including variation in longevity, skip or intermittent breeding, and persistent individual differences in reproductive success. Additional factors that can be important for some species are also discussed, including mating systems, population structure, sex reversal, reproductive compensation, captive propagation, and delayed maturity.

Partitioning variance in reproductive success, within years and across lifetimes.

Variance in reproductive success (, with k = number of offspring) plays a large role in determining the rate of genetic drift and the scope within which selection acts. Various frameworks have been proposed to parse factors that contribute to , but none has focused on age-specific values of , which indicate the degree to which reproductive skew is overdispersed (compared to the random Poisson expectation) among individuals of the same age and sex. Instead, within-age effects are generally lumped with residual variance and treated as "noise." Here, an ANOVA sums-of-squares framework is used to partition variance in annual and lifetime reproductive success into between-group and within-group components. For annual reproduction, the between-age effect depends on age-specific fecundity (b x), but relatively few empirical data are available on the within-age effect, which depends on ϕ x. By defining groups by age-at-death rather than age, the same ANOVA framework can be used to partition variance in lifetime reproductive success (LRS) into between-group and within-group components. Analytical methods are used to develop null-model expectations for random contributions to within-group and between-group components. For analysis of LRS, random variation in longevity appears as part of the between-group variance, and effects (if any) of skip breeding and persistent individual differences contribute to the within-group variance. Simulations are used to show that the methods for variance partitioning are asymptotically unbiased. Practical application is illustrated with empirical data for annual reproduction in American black bears and lifetime reproduction in Dutch great tits. Results show that overdispersed within-age variance (1) dominates annual in both male and female black bears, (2) is the primary factor that reduces annual effective size to a fraction of the number of adults, and (3) represents most of the opportunity for selection. In contrast, about a quarter of the variance in LRSin great tits can be attributed to random variation in longevity, and most of the rest is due to modest differences in fecundity with age estimated for a single cohort of females. R code is provided that reads generic input files for annual and lifetime reproductive success and allows users to conduct variance partitioning with their own data.

Hurricanes affect diversification among individual life courses of a primate population.

Extreme climatic events may influence individual-level variability in phenotypes, survival and reproduction, and thereby drive the pace of evolution. Climate models predict increases in the frequency of intense hurricanes, but no study has measured their impact on individual life courses within animal populations. We used 45 years of demographic data of rhesus macaques to quantify the influence of major hurricanes on reproductive life courses using multiple metrics of dynamic heterogeneity accounting for life course variability and life-history trait variances. To reduce intraspecific competition, individuals may explore new reproductive stages during years of major hurricanes, resulting in higher temporal variation in reproductive trajectories. Alternatively, individuals may opt for a single optimal life-history strategy due to trade-offs between survival and reproduction. Our results show that heterogeneity in reproductive life courses increased by 4% during years of major hurricanes, despite a 2% reduction in the asymptotic growth rate due to an average decrease in mean fertility and survival by that is, shortened life courses and reduced reproductive output. In agreement with this, the population is expected to achieve stable population dynamics faster after being perturbed by a hurricane ( ; 95% CI: 1.488, 1.538), relative to ordinary years . Our work suggests that natural disasters force individuals into new demographic roles to potentially reduce competition during unfavourable environments where mean reproduction and survival are compromised. Variance in lifetime reproductive success and longevity are differently affected by hurricanes, and such variability is mostly driven by survival.

The contribution of extra-pair paternity to the variation in lifetime and age-specific male reproductive success in a socially monogamous species.

In socially monogamous species, extra‐pair paternity (EPP) is predicted to increase variance in male reproductive success (RS) beyond that resulting from genetic monogamy, thus, increasing the “opportunity for selection” (maximum strength of selection that can act on traits). This prediction is challenging to investigate in wild populations because lifetime reproduction data are often incomplete. Moreover, age‐specific variances in reproduction have been rarely quantified. We analyzed 21 years of near‐complete social and genetic reproduction data from an insular population of Seychelles warblers (Acrocephalus sechellensis). We quantified EPP's contribution to lifetime and age‐specific opportunities for selection in males. We compared the variance in male genetic RS vs social (“apparent”) RS (RSap) to assess if EPP increased the opportunity for selection over that resulting from genetic monogamy. Despite not causing a statistically significant excess (19%) of the former over the latter, EPP contributed substantially (27%) to the variance in lifetime RS, similarly to within‐pair paternity (WPP, 39%) and to the positive WPP‐EPP covariance (34%). Partitioning the opportunity for selection into age‐specific (co)variance components, showed that EPP also provided a substantial contribution at most ages, varying with age. Therefore, despite possibly not playing the main role in shaping sexual selection in Seychelles warblers, EPP provided a substantial contribution to the lifetime and age‐specific opportunity for selection, which can influence evolutionary processes in age‐structured populations.

Longer telomeres during early life predict higher lifetime reproductive success in females but not males.

The mechanisms that contribute to variation in lifetime reproductive success are not well understood. One possibility is that telomeres, conserved DNA sequences at chromosome ends that often shorten with age and stress exposures, may reflect differences in vital processes or influence fitness. Telomere length often predicts longevity, but longevity is only one component of fitness and little is known about how lifetime reproductive success is related to telomere dynamics in wild populations. We examined the relationships between telomere length beginning in early life, telomere loss into adulthood and lifetime reproductive success in free-living house sparrows (Passer domesticus). We found that females, but not males, with longer telomeres during early life had higher lifetime reproductive success, owing to associations with longevity and not reproduction per year or attempt. Telomeres decreased with age in both sexes, but telomere loss was not associated with lifetime reproductive success. In this species, telomeres may reflect differences in quality or condition rather than the pace of life, but only in females. Sexually discordant selection on telomeres is expected to influence the stability and maintenance of within population variation in telomere dynamics and suggests that any role telomeres play in mediating life-history trade-offs may be sex specific.

Relative contributions of fixed and dynamic heterogeneity to variation in lifetime reproductive success in kestrels (Falco tinnunculus)

AbstractMany species show large variation in lifetime reproductive success (LRS), with a few individuals producing the majority of offspring. This variation can be explained by factors related to individuals (fixed heterogeneity) and stochastic differences in survival and reproduction (dynamic heterogeneity). In this study, we study the relative effects of these processes on the LRS of a Dutch Kestrel population, using three different methods. First, we extended neutral simulations by simulating LRS distributions of populations consisting of groups with increasingly different population parameters. Decomposition of total LRS variance into contributions from fixed and dynamic heterogeneity revealed that the proportion of fixed heterogeneity is probably lower than 10% of the total variance. Secondly, we used sensitivities of the mean and variance in LRS to each parameter to analytically show that it is impossible to get equal contributions of fixed and dynamic heterogeneity when only one parameter differs between groups. Finally, we computed the LRS probability distribution to show that even when all individuals have identical survival and reproduction rates, the variance in LRS is large (females: 27.52, males: 12.99). Although each method has its limitations, they all lead to the conclusion that the majority of the variation in kestrel LRS is caused by dynamic heterogeneity. This large effect of dynamic heterogeneity on LRS is similar to results for other species and contributes to the evidence that in most species the majority of individual variation in LRS is due to dynamic heterogeneity.

Variance in lifetime reproductive success of male polar bears

Abstract Despite the important role that population density plays in ecological and evolutionary processes, studies of solitary species that occur at low densities remain scarce. In the context of mating systems, density is expected to influence the ability of males to find and monopolize mates, in turn, influencing variance in lifetime mating/reproductive success and the opportunity for selection. Herein, we investigate variance in male lifetime mating success (LMS), lifetime reproductive success (LRS), and the mating system of a sexually dimorphic carnivore that occurs at low densities, the polar bear (Ursus maritimus). Across 17 cohorts, born from 1975 to 1991, male LMS ranged from 0 to10 mates and LRS from 0 to 14 cubs; 40% of known-age males were not known to have reproduced. The opportunity for sexual selection (Is = 1.66, range = 0.60–4.99) and selection (I = 1.76, range: 0.65–4.89) were low compared to species with similar levels of sexual size dimorphism. Skew in male LRS was also low but significant for most cohorts indicating nonrandom reproductive success. Age-specific reproductive success was biased toward males from 11 to 17 years of age, with variation in fecundity (54%) but not longevity (10%) playing an important role in male reproduction. Our results support a growing body of evidence that suggests that male-biased size dimorphism and polygynous mating systems need not be associated with high variance in male mating and/or reproductive success.

Lifetime reproductive success of northern elephant seals (Mirounga angustirostris)

Lifetime reproductive success of individuals in a natural population provides an estimate of Darwinian fitness. We calculated lifetime reproductive success in a colony of female northern elephant seals (Mirounga angustirostris (Gill, 1866)) by monitoring annual breeding throughout life of 7735 female weanlings marked individually at Año Nuevo, California, USA, from 1963 to 2005. Great variation in lifetime reproductive success was evident in three aspects of life history: (1) 75% of the females died before reaching breeding age and produced no pups; (2) nearly half of the survivors bred for only a few years before dying, and young females had low weaning success; (3) less than 1% of the females in the sample were exceptionally successful producing up to 20 pups in life. Many females that bred early, while still growing, had decreased lifespan, low weaning success, and lower lifetime reproductive success than females that postponed first breeding. Exceptional reproductive success was associated with giving birth annually, living long (up to age 23), and weaning large pups that were more likely to survive and breed. We conclude that there is strong selection for increased lifespan and multiparous supermoms that contribute significantly to pup production in the next generation.

Male body size, dominance rank and strategic use of aggression in a group-living mammal

Body size is a key determinant of male fighting ability and reproductive success in many animal species, but relationships between these variables have only rarely been examined in group-living animals in which body size often correlates with dominance rank. We examined the relationships between body size (crest height, back breadth and body length), dominance rank, alpha male tenure length, number of adult females and patterns of aggression in 26 wild adult male mountain gorillas, Gorilla beringei beringei, living in multimale groups. A composite measure combining crest height and back breadth (variables were highly correlated and combined into a crest–back score), but not body length, significantly correlated with dominance rank, alpha male tenure length and number of adult females per group. The alpha male had the largest crest–back score in six of the seven groups, and in the majority of dyads the male with the higher crest–back score was higher ranking. The frequency (and intensity on mating days) of aggressive contests was higher between males close in rank. Additionally, aggression occurred more frequently when the initiator was larger than the recipient. Our results suggest that factors other than body size are likely to influence dominance rank, but large size helps males attain and retain high dominance rank, probably leading to greater reproductive success. Further studies on how the timing and intensity of male–male competition influences life history trade-offs between investment in secondary sexual characteristics, body condition and survival may explain variance in lifetime reproductive success within and between species.

Extrapair mating and the strength of sexual selection: insights from a polymorphic species.

Extrapair mating could drive sexual selection in socially monogamous species, but support for this hypothesis remains equivocal. We used lifetime fitness data and a unique model species, the dimorphic white-throated sparrow (Zonotrichia albicollis), to examine how extrapair mating affects the potential for sexual selection. In this species, the morphs employ distinct reproductive strategies, with white males pursuing extrapair mating at higher rates than tan counterparts. Social and extrapair mating is disassortative by morph, with paternity exchange occurring primarily between pairs composed of white males and tan females. Bateman gradients and Jones indexes indicated stronger sexual selection via mate numbers in white males than in females and tan males, and generally did not differ between females as compared with tan males. Extrapair mating contributed more to the Bateman gradient for white than tan males, and white males also had higher variance in annual reproductive success. However, variance in lifetime reproductive success did not differ between morphs or sexes. Moreover, extrapair mating did not increase variance in male reproductive success relative to apparent patterns, and within-pair success accounted for much more variance than extrapair success. Thus, extrapair mating by white males increases Bateman gradients and the potential for sexual selection via mate numbers. However, our latter results support previous research suggesting that extrapair mating may play a limited role in driving the overall potential for sexual selection.

Revealing life-history traits by contrasting genetic estimations with predictions of effective population size.

Effective population size, a central concept in conservation biology, is now routinely estimated from genetic surveys and can also be theoretically predicted from demographic, life-history, and mating-system data. By evaluating the consistency of theoretical predictions with empirically estimated effective size, insights can be gained regarding life-history characteristics and the relative impact of different life-history traits on genetic drift. These insights can be used to design and inform management strategies aimed at increasing effective population size. We demonstrated this approach by addressing the conservation of a reintroduced population of Asiatic wild ass (Equus hemionus). We estimated the variance effective size (Nev ) from genetic data (N ev =24.3) and formulated predictions for the impacts on Nev of demography, polygyny, female variance in lifetime reproductive success (RS), and heritability of female RS. By contrasting the genetic estimation with theoretical predictions, we found that polygyny was the strongest factor affecting genetic drift because only when accounting for polygyny were predictions consistent with the genetically measured Nev . The comparison of effective-size estimation and predictions indicated that 10.6% of the males mated per generation when heritability of female RS was unaccounted for (polygyny responsible for 81% decrease in Nev ) and 19.5% mated when female RS was accounted for (polygyny responsible for 67% decrease in Nev ). Heritability of female RS also affected Nev ; hf2=0.91 (heritability responsible for 41% decrease in Nev ). The low effective size is of concern, and we suggest that management actions focus on factors identified as strongly affecting Nev, namely, increasing the availability of artificial water sources to increase number of dominant males contributing to the gene pool. This approach, evaluating life-history hypotheses in light of their impact on effective population size, and contrasting predictions with genetic measurements, is a general, applicable strategy that can be used to inform conservation practice.

Pluck or Luck: Does Trait Variation or Chance Drive Variation in Lifetime Reproductive Success?

While there has been extensive interest in how intraspecific trait variation affects ecological processes, outcomes are highly variable even when individuals are identical: some are lucky, while others are not. Trait variation is therefore important only if it adds substantially to the variability produced by luck. We ask when trait variation has a substantial effect on variability in lifetime reproductive success (LRS), using two approaches: (1) we partition the variation in LRS into contributions from luck and trait variation and (2) we ask what can be inferred about an individual's traits and with what certainty, given their observed LRS. In theoretical stage- and size-structured models and two empirical case studies, we find that luck usually dominates the variance of LRS. Even when individuals differ substantially in ways that affect expected LRS, unless the effects of luck are substantially reduced (e.g., low variability in reproductive life span or annual fecundity), most variance in lifetime outcomes is due to luck, implying that departures from "null" models omitting trait variation will be hard to detect. Luck also obscures the relationship between realized LRS and individual traits. While trait variation may influence the fate of populations, luck often governs the lives of individuals.

Life expectancy, maximum longevity and lifetime reproductive success in female Thornicroft's giraffe in Zambia

AbstractGestation and longevity scale with body mass across taxa, yet within size dimorphic taxa, males tend to have reduced lifespans compared with females. Testing life history models, and accounting for sex differences in longevity, requires obtaining accurate longitudinal data from wild populations. We provide the first report describing key life history parameters from a long‐term study of giraffes in Africa. We followed a population of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti) in Zambia for over 40 years. Maximum longevity among females was approximately 28 years, with lifespan accounting for 81% of the variance in lifetime reproductive success. Average adult female life expectancy was no different than average adult male life expectancy. However, the breeding lifespan of males was about half that of females, while maximum lifespan of males was 75% that of females. Our findings support the suggestion that sex differences in maximum lifespan arise from stronger selection for lengthy lives in females than in males. Among females, longer lives are associated with greater reproductive output.

USE OF AGE‐ AND STAGE‐STRUCTURED MATRIX MODELS TO PREDICT LIFE HISTORY SCHEDULES FOR SEMELPAROUS POPULATIONS

AbstractMany studies of semelparous salmon populations use Leslie matrices that classify individuals on the basis of age alone and do not explicitly impose death upon reproduction. Although these models may suffice for studying long‐term population dynamics (like asymptotic growth rate), they do not accurately represent the diversity of individual life history outcomes in semelparous populations. Cohorts breeding at different ages have different life history traits (e.g., age at first reproduction and remaining life expectancy) that are obscured in Leslie models and this distorts our understanding of life history diversity and its importance for semelparous population dynamics. We present a simple transformation that uses age‐specific breeding probabilities to reconfigure Leslie matrices as explicitly semelparous models. Explicitly semelparous models conserve asymptotic measures like population growth rate, vital rate elasticities, life expectancy at birth, and generation time but also better predict life history schedules and reproductive values. Strictly age‐classified Leslie models underestimate ages at first reproduction and mean ages at death for older breeders but overestimate mean ages at death for early breeders. Leslie models also slightly overestimate variance in lifetime reproductive success, and underestimate entropy exhibited by life history outcomes.

The effects of asymmetric competition on the life history of Trinidadian guppies.

The effects of asymmetric interactions on population dynamics has been widely investigated, but there has been little work aimed at understanding how life history parameters like generation time, life expectancy and the variance in lifetime reproductive success are impacted by different types of competition. We develop a new framework for incorporating trait‐mediated density‐dependence into size‐structured models and use Trinidadian guppies to show how different types of competitive interactions impact life history parameters. Our results show the degree of symmetry in competitive interactions can have dramatic effects on the speed of the life history. For some vital rates, shifting the competitive superiority from small to large individuals resulted in a doubling of the generation time. Such large influences of competitive symmetry on the timescale of demographic processes, and hence evolution, highlights the interwoven nature of ecological and evolutionary processes and the importance of density‐dependence in understanding eco‐evolutionary dynamics.

Lifetime reproductive success of Snowy Plovers in coastal northern California

ABSTRACT Conserving threatened and endangered species requires knowledge of breeding productivity and factors that cause variation in reproductive success. We summarized 13 years of lifetime reproductive success (LRS) data for 195 individually marked Snowy Plovers (Charadrius nivosus) breeding in Humboldt County, CA. Reproductive success was highly skewed among individuals with 13% of individuals (nmales = 12, nfemales = 14) producing ~50% of fledglings; by contrast, 71% (n = 64) of males and 72% (n = 76) of females produced 2 or fewer during their lifetime. Variance in LRS was best explained by substrate (~100% of Akaike weight), with plovers breeding on gravel having significantly higher LRS compared to those on sandy substrates. Other measures of habitat quality, including use of nest exclosures, as well as corvid and human activity, were not significant predictors of LRS. Results indicated that enhancing the cryptic nature of substrates (for eggs and chicks) may be a productive means of increasing rep...

Very low levels of direct additive genetic variance in fitness and fitness components in a red squirrel population.

A trait must genetically correlate with fitness in order to evolve in response to natural selection, but theory suggests that strong directional selection should erode additive genetic variance in fitness and limit future evolutionary potential. Balancing selection has been proposed as a mechanism that could maintain genetic variance if fitness components trade off with one another and has been invoked to account for empirical observations of higher levels of additive genetic variance in fitness components than would be expected from mutation–selection balance. Here, we used a long-term study of an individually marked population of North American red squirrels (Tamiasciurus hudsonicus) to look for evidence of (1) additive genetic variance in lifetime reproductive success and (2) fitness trade-offs between fitness components, such as male and female fitness or fitness in high- and low-resource environments. “Animal model” analyses of a multigenerational pedigree revealed modest maternal effects on fitness, but very low levels of additive genetic variance in lifetime reproductive success overall as well as fitness measures within each sex and environment. It therefore appears that there are very low levels of direct genetic variance in fitness and fitness components in red squirrels to facilitate contemporary adaptation in this population.

Intra-sexual selection in cooperative mammals and birds: why are females not bigger and better armed?

In cooperatively breeding mammals and birds, intra-sexual reproductive competition among females may often render variance in reproductive success higher among females than males, leading to the prediction that intra-sexual selection in such species may have yielded the differential exaggeration of competitive traits among females. However, evidence to date suggests that female-biased reproductive variance in such species is rarely accompanied by female-biased sexual dimorphisms. We illustrate the problem with data from wild Damaraland mole-rat, Fukomys damarensis, societies: the variance in lifetime reproductive success among females appears to be higher than that among males, yet males grow faster, are much heavier as adults and sport larger skulls and incisors (the weapons used for fighting) for their body lengths than females, suggesting that intra-sexual selection has nevertheless acted more strongly on the competitive traits of males. We then consider potentially general mechanisms that could explain these disparities by tempering the relative intensity of selection for competitive trait exaggeration among females in cooperative breeders. Key among these may be interactions with kin selection that could nevertheless render the variance in inclusive fitness lower among females than males, and fundamental aspects of the reproductive biology of females that may leave reproductive conflict among females more readily resolved without overt physical contests.

Decomposing variation in male reproductive success: age‐specific variances and covariances through extra‐pair and within‐pair reproduction

Age-specific variances and covariances in reproductive success shape the total variance in lifetime reproductive success (LRS), age-specific opportunities for selection, and population demographic variance and effective size. Age-specific (co)variances in reproductive success achieved through different reproductive routes must therefore be quantified to predict population, phenotypic and evolutionary dynamics in age-structured populations. While numerous studies have quantified age-specific variation in mean reproductive success, age-specific variances and covariances in reproductive success, and the contributions of different reproductive routes to these (co)variances, have not been comprehensively quantified in natural populations. We applied 'additive' and 'independent' methods of variance decomposition to complete data describing apparent (social) and realised (genetic) age-specific reproductive success across 11 cohorts of socially monogamous but genetically polygynandrous song sparrows (Melospiza melodia). We thereby quantified age-specific (co)variances in male within-pair and extra-pair reproductive success (WPRS and EPRS) and the contributions of these (co)variances to the total variances in age-specific reproductive success and LRS. 'Additive' decomposition showed that within-age and among-age (co)variances in WPRS across males aged 2-4 years contributed most to the total variance in LRS. Age-specific (co)variances in EPRS contributed relatively little. However, extra-pair reproduction altered age-specific variances in reproductive success relative to the social mating system, and hence altered the relative contributions of age-specific reproductive success to the total variance in LRS. 'Independent' decomposition showed that the (co)variances in age-specific WPRS, EPRS and total reproductive success, and the resulting opportunities for selection, varied substantially across males that survived to each age. Furthermore, extra-pair reproduction increased the variance in age-specific reproductive success relative to the social mating system to a degree that increased across successive age classes. This comprehensive decomposition of the total variances in age-specific reproductive success and LRS into age-specific (co)variances attributable to two reproductive routes showed that within-age and among-age covariances contributed substantially to the total variance and that extra-pair reproduction can alter the (co)variance structure of age-specific reproductive success. Such covariances and impacts should consequently be integrated into theoretical assessments of demographic and evolutionary processes in age-structured populations.

Reproductive success of female leopards Panthera pardus: the importance of top‐down processes

Abstract Long‐term studies on large felids are rare and yet they yield data essential to understanding the behaviour of species and the factors that facilitate their conservation. We used the most extensive data set so far compiled on leopards Panthera pardus to establish baseline reproductive parameters for females and to determine the demographic and environmental factors that affect their lifetime reproductive success. We used comprehensive sightings reports and photographs from ecotourism lodges in the Sabi Sand Game Reserve, South Africa, to reconstruct life histories for 44 female leopards that gave birth to 172 litters over a 32‐year period. Leopards appeared to exhibit a birth pulse; most litters were born in the wet season, particularly in December. Mean age at first parturition (n = 26, mean ± standard error = 46 ± 2 months, range = 33–62) was older than previously recorded, possibly due to elevated intraspecific competition. Average litter size was 1.9 ± 0.1 (n = 140, range = 1–3) and declined with maternal age. Age of litters at independence (n = 52, 19 ± 1 months, range = 9–31) was inversely related to prey abundance but did not affect the likelihood of recruitment of offspring. Interbirth intervals differed following successful litters (in which at least one cub survived to independence; n = 55, 25 ± 1 months, range = 14–39) and unsuccessful litters (n = 46, 11 ± 1 months, range = 4–36), as did the time taken to replace litters. Variation in lifetime reproductive success was influenced mainly by differences in cub survival, which was related to maternal age and vulnerability to infanticide. Cub survival (37%) declined as females got older, perhaps because mothers relinquished portions of their home ranges to philopatric daughters. Male leopards were responsible for many (40%) cub deaths and females appeared to adopt severalstrategies to counter the risk of infanticide, including paternity confusion and displaying a period of reduced fertility immediately after a resident male was replaced. Our results suggest that the reproductive success of female leopards is regulated primarily by top‐down processes. This should be taken into account in management decisions, particularly when managers are considering the implementation of invasive activities such as legal trophy hunting.