Lifetime Reproductive Success Research Articles

Modeling of senescent cell dynamics predicts a late-life decrease in cancer incidence.

Current oncogenic theories state that tumors arise from cell lineages that sequentially accumulate (epi)mutations, progressively turning healthy cells into carcinogenic ones. While those models found some empirical support, they are little predictive of intraspecies age-specific cancer incidence and of interspecies cancer prevalence. Notably, in humans and lab rodents, a deceleration (and sometimes decline) of cancer incidence rate has been found at old ages. Additionally, dominant theoretical models of oncogenesis predict that cancer risk should increase in large and/or long-lived species, which is not supported by empirical data. Here, we explore the hypothesis that cellular senescence could explain those incongruent empirical patterns. More precisely, we hypothesize that there is a trade-off between dying of cancer and of (other) ageing-related causes. This trade-off between organismal mortality components would be mediated, at the cellular scale, by the accumulation of senescent cells. In this framework, damaged cells can either undergo apoptosis or enter senescence. Apoptotic cells lead to compensatory proliferation, associated with an excess risk of cancer, whereas senescent cell accumulation leads to ageing-related mortality. To test our framework, we build a deterministic model that first describes how cells get damaged, undergo apoptosis, or enter senescence. We then translate those cellular dynamics into a compound organismal survival metric also integrating life-history traits. We address four different questions linked to our framework: can cellular senescence be adaptive, do the predictions of our model reflect epidemiological patterns observed among mammal species, what is the effect of species sizes on those answers, and what happens when senescent cells are removed? Importantly, we find that cellular senescence can optimize lifetime reproductive success. Moreover, we find that life-history traits play an important role in shaping the cellular trade-offs. Overall, we demonstrate that integrating cellular biology knowledge with eco-evolutionary principles is crucial to solve parts of the cancer puzzle.

Evidence of an Absence of Inbreeding Depression in a Wild Population of Weddell Seals (Leptonychotes weddellii).

Inbreeding depression can reduce the viability of wild populations. Detecting inbreeding depression in the wild is difficult; developing accurate estimates of inbreeding can be time and labor intensive. In this study, we used a two-step modeling procedure to incorporate uncertainty inherent in estimating individual inbreeding coefficients from multilocus genotypes into estimates of inbreeding depression in a population of Weddell seals (Leptonychotes weddellii). The two-step modeling procedure presented in this paper provides a method for estimating the magnitude of a known source of error, which is assumed absent in classic regression models, and incorporating this error into inferences about inbreeding depression. The method is essentially an errors-in-variables regression with non-normal errors in both the dependent and independent variables. These models, therefore, allow for a better evaluation of the uncertainty surrounding the biological importance of inbreeding depression in non-pedigreed wild populations. For this study we genotyped 154 adult female seals from the population in Erebus Bay, Antarctica, at 29 microsatellite loci, 12 of which are novel. We used a statistical evidence approach to inference rather than hypothesis testing because the discovery of both low and high levels of inbreeding are of scientific interest. We found evidence for an absence of inbreeding depression in lifetime reproductive success, adult survival, age at maturity, and the reproductive interval of female seals in this population.

Oviposition experience promotes active reproductive behaviour in a synovigenic parasitoid

Parasitoids are important insects that are commonly released into the environment to reduce the population sizes of pest species. The lifetime reproductive success of parasitoids mainly depends on host availability and the availability of mature eggs. Consequently, it is predicted that female wasps must balance the risk of egg or host (time) limitation with maximized lifetime fecundity. Typically, synovigenic females, which continue to mature eggs throughout their lifetime, have been shown to adjust their egg production rate in response to environmental variations in host availability to reduce the risk of egg limitation. In this study, we found that in a synovigenic egg parasitoid, Anastatus japonicus (Hymenoptera: Eupelmidae), the oviposition experience of Ana. japonicus females significantly enhanced the egg load and increased the rate of mature egg production. However, in contrast to other studies, the experience of contact with a host did not significantly affect the egg load in females. This result suggests that the overall oviposition experience might induce an adjustment and accelerate egg maturation in Ana. japonicus and is likely more important in egg maturation than transitory host contract. In addition to affecting the egg load, oviposition experience influenced Ana. japonicus female reproductive behaviour, which shifted virgin female behavioural preferences from mating to oviposition and laying more eggs per clutch. Our study provides an optimal strategy for the post-oviposition release of Ana. japonicus, an egg parasitoid of several lepidopteran forest pests, to improve biocontrol effectiveness.

Genomic signatures of inbreeding depression for a threatened Aotearoa New Zealand passerine.

For small and isolated populations, the increased chance of mating between related individuals can result in a substantial reduction in individual and population fitness. Despite the increasing availability of genomic data to measure inbreeding accurately across the genome, inbreeding depression studies for threatened species are still scarce due to the difficulty of measuring fitness in the wild. Here, we investigate inbreeding and inbreeding depression for the extensively monitored Tiritiri Mātangi island population of a threatened Aotearoa New Zealand passerine, the hihi (Notiomystis cincta). First, using a custom 45 k single nucleotide polymorphism (SNP) array, we explore genomic inbreeding patterns by inferring homozygous segments across the genome. Although all individuals have similar levels of ancient inbreeding, highly inbred individuals are affected by recent inbreeding, which can probably be explained by bottleneck effects such as habitat loss after European arrival and their translocation to the island in the 1990s. Second, we investigate genomic inbreeding effects on fitness, measured as lifetime reproductive success, and its three components, juvenile survival, adult annual survival and annual reproductive success, in 363 hihi. We find that global inbreeding significantly affects juvenile survival but none of the remaining fitness traits. Finally, we employ a genome-wide association approach to test the locus-specific effects of inbreeding on fitness, and identify 13 SNPs significantly associated with lifetime reproductive success. Our findings suggest that inbreeding depression does impact hihi, but at different genomic scales for different traits, and that purging has therefore failed to remove all variants with deleterious effects from this population of conservation concern.

Active breeding seabirds prospect alternative breeding colonies

Compared to other animal movements, prospecting by adult individuals for a future breeding site is commonly overlooked. Prospecting influences the decision of where to breed and has consequences on fitness and lifetime reproductive success. By analysing movements of 31 satellite- and GPS-tracked gull and tern populations belonging to 14 species in Europe and North America, we examined the occurrence and factors explaining prospecting by actively breeding birds. Prospecting in active breeders occurred in 85.7% of studied species, across 61.3% of sampled populations. Prospecting was more common in populations with frequent inter-annual changes of breeding sites and among females. These results contradict theoretical models which predict that prospecting is expected to evolve in relatively predictable and stable environments. More long-term tracking studies are needed to identify factors affecting patterns of prospecting in different environments and understand the consequences of prospecting on fitness at the individual and population level.

Unveiling demographic and mating strategies of Panthera onca in the Pantanal, Brazil.

We conducted the first long-term and large-scale study of demographic characteristics and reproductive behavior in a wild jaguar (Panthera onca) population. Data were collected through a combination of direct observations and camera trapping on a study area that operates both as a cattle ranch and ecotourism destination. Jaguars exhibited two birth peaks: April/May and October/November, that are the end and the beginning of the wet season in the Pantanal, respectively. The average litter size was 1.43 ± 0.65. Single cubs made up a total of 65.7% of the births, and we found a slight predominance of females (1.15:1 ratio) in litters. The mean age at independence was 17.6 ± 0.98 months, with sex-biased dispersal, with all males (n = 27) leaving the natal home range and 63.6% of females exhibiting philopatry. The interbirth intervals were 21.8 ± 3.2 months and the mean age at first parturition was 31.8 ± 4.2 months. Our results estimated a lifetime reproductive success for female jaguars of 8.13 cubs. Our observations also indicate that female jaguars can display mating behavior during cub rearing or pregnancy, representing 41.4% of the consorts and copulations recorded. We speculate that this behavior has evolved as a defense against infanticide and physical harm to the female. To our knowledge, this is the first time that such behavior is described for this species. All aggressive interactions between females involved the presence of cubs, following the offspring-defense hypothesis, that lead to territoriality among females in mammals, regardless of food availability. In the face of growing threats to this apex predator, this work unveils several aspects of its natural history, representing a baseline for comparison with future research and providing critical information for population viability analysis and conservation planning in the long term.

Изучение биологии гнездования беркута с помощью фотоловушек

Trap camera was installed in an active nest of Golden Eagles (Aquila chrysaetos) in Eastern Rhodopes, Bulgaria. The recordings encompassed 45 days in June and July and the camera took 12 900 photos, from the time when the single nestling was ca. 32 days old until its fledging. 84 food deliveries were recorded, with the prey identified in 76 of them. Land tortoises dominated the diet (65% of the items), followed by Edible Dormouse (Glis glis) (10.5%), snakes (7.9%), and European Hare (Lepus europaeus) (5.2%). The two adults were easily individually identifiable by plumage characteristics, especially on the medium coverts. We recorded unexpectedly high intensity of adding the green material to the nest and cleaning the nest from prey remains, 105 and 35 cases respectively, both done almost exclusively by the female, while the male delivered most of the food – 60 vs 21 of the cases. The female was roosting in the nest with the nestling until when it was 50 days old, after which roosted in the nest only once during a rainy night when nestling was 58 days old. The method provided opportunity for following the lifetime reproductive success of unmarked Golden Eagles.

Reproductive performance and sex ratio adjustment of the wild boar (Sus scrofa) in South Korea

The wild boar (Sus scrofa), a polygynous species, rapidly expanded its geographical range and increased its population size in South Korea following the extinction of large carnivores and changes to rural environments. Understanding wild boar reproductive traits and strategies is essential for their effective management; however, studies in this area are lacking. Using samples collected from hunting bags, the relationships between 1) litter size and female weight and 2) fetal sex ratio and female body condition were examined to understand wild boar life-history strategies. Wild boars showed a seasonal breeding pattern that maximized reproduction. Litter size (mean = 5.7 ± 1.7) was correlated with female weight, whereas fetal sex ratio was not explained by female body condition. However, the heaviest ranked fetuses within the litters were male-biased. Wild boars aged three years or less accounted for 90% of the total population, and sexual dimorphism developed from two years of age. Considering that their reproductive strategy is more effective (i.e., early gestation and large litter size) than that of other polygynous species, the Trivers–Willard model was not supported for the wild boars in this study. Instead, females adjusted the sex of the heaviest fetus in the litter to maximize lifetime reproductive success.

Sex-dependent effects of parental age on offspring fitness in a cooperatively breeding bird.

Parental age can have considerable effects on offspring phenotypes and health. However, intergenerational effects may also have longer term effects on offspring fitness. Few studies have investigated parental age effects on offspring fitness in natural populations while also testing for sex- and environment-specific effects. Further, longitudinal parental age effects may be masked by population-level processes such as the selective disappearance of poor-quality individuals. Here, we used multigenerational data collected on individually marked Seychelles warblers (Acrocephalus sechellensis) to investigate the impact of maternal and paternal age on offspring life span and lifetime reproductive success. We found negative effects of maternal age on female offspring life span and lifetime reproductive success, which were driven by within-mother effects. There was no difference in annual reproductive output of females born to older versus younger mothers, suggesting that the differences in offspring lifetime reproductive success were driven by effects on offspring life span. In contrast, there was no association between paternal age and female offspring life span or either maternal or paternal age and male offspring life span. Lifetime reproductive success, but not annual reproductive success, of male offspring increased with maternal age, but this was driven by between-mother effects. No paternal age effects were found on female offspring lifetime reproductive success but there was a positive between-father effect on male offspring lifetime reproductive success. We did not find strong evidence for environment-dependent parental age effects. Our study provides evidence for parental age effects on the lifetime fitness of offspring and shows that such effects can be sex dependent. These results add to the growing literature indicating the importance of intergenerational effects on long-term offspring performance and highlight that these effects can be an important driver of variation in longevity and fitness in the wild.

Different proxies, different stories? Imperfect correlations and different determinants of fitness in bighorn sheep

Measuring individual fitness empirically is required to assess selective pressures and predicts evolutionary changes in nature. There is, however, little consensus on how fitness should be empirically estimated. As fitness proxies vary in their underlying assumptions, their relative sensitivity to individual, environmental, and demographic factors may also vary. Here, using a long-term study, we aimed at identifying the determinants of individual fitness in bighorn sheep (Ovis canadensis) using seven fitness proxies. Specifically, we compared four-lifetime fitness proxies: lifetime breeding success, lifetime reproductive success, individual growth rate, individual contribution to population growth, and three multi-generational proxies: number of granddaughters, individual descendance in the next generation, and relative genetic contribution to the next generation. We found that all proxies were positively correlated, but the magnitude of the correlations varied substantially. Longevity was the main determinant of most fitness proxies. Individual fitness calculated over more than one generation was also affected by population density and growth rate. Because they are affected by contrasting factors, our study suggests that different fitness proxies should not be used interchangeably as they may convey different information about selective pressures and lead to divergent evolutionary predictions. Uncovering the mechanisms underlying variation in individual fitness and improving our ability to predict evolutionary change might require the use of several, rather than one, the proxy of individual fitness.

How do helpers slow senescence in a Tibetan cooperatively breeding bird?

AbstractSociality is known to be capable of slowing individual senescence, but it is unclear whether the effect differs for reproduction versus survival in a sex‐specific manner. Here we predict that social benefits are directed toward (1) somatic maintenance in harsh environments where high survival prospects of adults over young intensify the trade‐off between current and future reproduction, and (2) females that invest more in reproduction and have a greater marginal effect if their survival is improved by reducing the cost of reproduction. These two predictions are tested with cooperatively breeding Tibetan ground tits (Pseudopodoces humilis). Across the lifetime, both mothers and fathers with helpers did not differ in brood size at fledging from their counterparts without helpers. The presence of helpers reduced survival senescence of both parents, but mothers benefited more than fathers from receiving help. Consequently, the inherent sex difference in life span and along with lifetime reproductive success, as expressed in breeders of never‐receiving help, became no longer obvious. The model of social modulation for senescence should facilitate the persistence of cooperative breeding in challenging environments.

Combined effects of exploitation and environmental change on life history: a comparative analysis on Atlantic herring

Abstract The consequences of fisheries-induced evolution on stock productivity and yield depend, to a large extent, on the general prospects for growth and survival. Here, we compare the selection pressures imposed by two distinct patterns of exploitation—principally targeting spawning or non-spawning aggregations—on age at maturity among 15 Canadian stocks of Atlantic herring (Clupea harengus) that have exhibited a consistent pattern of length-at-age responses to common large-scale environmental drivers since the 1960s. In accordance with expectations for maturity-dependent harvesting, the establishment of a spawner-targeted fishery in the southern Gulf of St. Lawrence coincided with a shift towards delayed maturity in both resident stocks, whereas stocks elsewhere subject to fisheries that also exploited juveniles more commonly exhibited trends towards earlier maturity. Despite these differences, we find that environmentally driven changes in length at maturation, combined with total mortality, may overwhelmingly determine lifetime reproductive success and possibly fitness. By linking phenotypic changes experienced in the juvenile period to simple correlates of egg production in mature age classes, our study highlights the importance of managing fisheries in the context of ubiquitous but contrasting environmental constraints on life histories.

Stochasticity directs adaptive evolution toward nonequilibrium evolutionary attractors.

Stochastic processes such as genetic drift may hinder adaptation, but the effect of such stochasticity on evolution via its effect on ecological dynamics is poorly understood. Here we evaluate patterns of adaptation in a population subject to variation in demographic stochasticity. We show that stochasticity can alter population dynamics and lead to evolutionary outcomes that are not predicted by classic eco-evolutionary modeling approaches. We also show, however, that these outcomes are governed by nonequilibrium evolutionary attractors-these are maxima in lifetime reproductive success when stochasticity keeps the ecological system away from the deterministic equilibrium. These NEEAs alter the path of evolution but are not visible through the equilibrium lens that underlies much evolutionary theory. Our results reveal that considering population processes during transient periods can greatly improve our understanding of the path and pace of evolution.

Reproductive skews of territorial species in heterogeneous landscapes

The socio–spatial organization of populations of territorial species refers to the way distance, variation in social status and variation in resource availability interact to shape the way territory‐bounded individuals are distributed. I investigated the resulting reproductive skew, i.e. the emergence of a few high performers amidst a majority of mediocre performers, using lifetime reproductive success data from 46 species living in contrasting landscape configurations. I applied three skew statistics that captured different aspects of the individual heterogeneity: the skewness, the Gini coefficient of inequalities and Hill's index of heavy‐tailed distribution. I also used a simplistic theoretical model, in which only immediate neighbors interacted. In the comparative analysis, the distribution of lifetime reproductive success was more skewed but also more egalitarian in homogeneous than heterogeneous landscapes, after controlling for the allometric effect of the body mass. The simulations indicated that a possible explanatory mechanism for the change in egalitarian properties is that some individuals hide behind their neighbors in the socio‐spatial organization. They therefore secure a better territory than if the distribution was despotic. The ideal despotic model without spatial restriction on interactions could not explain the correlations. This work emphasizes that the shape of the distribution of fitness components conveys information besides the mean and the variance, and the potential for nontrivial effects to emerge from simple movement heuristics.

Mating systems in birds

Most vertebrates, including all mammals and birds, are sexually reproducing organisms. The implication of sexual reproduction is that an adult bird exists because a male transferred sperm to the reproductive tract of a female during copulation, one of the sperm fertilized an egg, and the ensuing embryo developed into an individual male or female that in turn can reproduce. This fundamental feature of sexual life - the need to bring together male and female units (genomes, gametes, individuals) in a sequence of interactions - is both fascinating and fraught with conflict, because selection acts on individuals to maximize their own lifetime reproductive success (or on individual units of DNA to maximize the number of copies of themselves that remain in the gene pool). As a consequence, a bewildering diversity of reproductive traits and behaviours has evolved.

Evolutionary consequences of pesticide exposure include transgenerational plasticity and potential terminal investment transgenerational effects.

Transgenerational plasticity, the influence of the environment experienced by parents on the phenotype and fitness of subsequent generations, is being increasingly recognized. Human-altered environments, such as those resulting from the increasing use of pesticides, may be major drivers of such cross-generational influences, which in turn may have profound evolutionary and ecological repercussions. Most of these consequences are, however, unknown. Whether transgenerational plasticity elicited by pesticide exposure is common, and the consequences of its potential carryover effects on fitness and population dynamics, remains to be determined. Here, we investigate whether exposure of parents to a common pesticide elicits intra-, inter-, and transgenerational responses (in F0, F1, and F2 generations) in life history (fecundity, longevity, and lifetime reproductive success), in an insect model system, the seed beetle Callosobruchus maculatus. We also assessed sex specificity of the effects. We found sex-specific and hormetic intergenerational and transgenerational effects on longevity and lifetime reproductive success, manifested both in the form of maternal and paternal effects. In addition, the transgenerational effects via mothers detected in this study are consistent with a new concept: terminal investment transgenerational effects. Such effects could underlie cross-generational responses to environmental perturbation. Our results indicate that pesticide exposure leads to unanticipated effects on population dynamics and have far-reaching ecological and evolutionary implications.

Predator avoidance influences selection of neonatal lambing habitat by Sierra Nevada bighorn sheep

AbstractTo improve lifetime reproductive success, maternal ungulates should pursue behavioral strategies that reduce risk of offspring mortality. Predation is a leading cause of neonatal mortality; thus, maternal ungulates should select habitat that reduces risk of predation on vulnerable neonates. We examined selection of habitat across subpopulations of federally endangered Sierra Nevada bighorn sheep (Ovis canadensis sierrae) within Sierra Nevada, California, USA, 2008–2018. Our objectives were to understand how maternal Sierra Nevada bighorn minimize risk of predation through selection of habitat and to quantify current and future potential neonatal habitat across the Sierra Nevada. Mountain lions (Puma concolor) are the predominant predators of adult Sierra Nevada bighorn; thus, we hypothesized females would select habitat that minimized risk of predation from mountain lions. We developed a used‐available resource selection function to quantify patterns of habitat selection and produced predictive maps identifying highly selected lambing habitat across the Sierra Nevada. Our top model demonstrated females selected habitat where the relative probability of encountering mountain lions was low and near escape terrain. Selection for vegetation type was dependent on risk of encounter with mountain lions; when risk of encounter was low, females selected shrub cover. This suggests that when risk of encountering mountain lions was low, females may have selected areas with dense shrubs to reduce risk of being detected by other predators such as coyotes (Canis latrans) and golden eagles (Aquila chrysaetos). As risk of encountering mountain lions increased, females avoided all vegetation types other than barren, suggesting that despite increased risk of encounter, females may have been able to mitigate overall mortality risk by selecting habitat where approaching mountain lions could be detected and avoided. Our predictive models and resulting maps demonstrate differences in prevalence and connectivity of highly selected lambing habitat across subpopulations, which may explain differences in lamb recruitment and adequacy of lambing habitat observed among subpopulations. Recolonization into historical ranges and increasing connectivity between Sierra Nevada bighorn subpopulations is an important conservation need for species recovery and long‐term viability of fragmented subpopulations.

The Feast and the Famine: Spring Body Mass Variations and Life History Traits in a Pulse Resource Ecosystem.

AbstractPhenotypic plasticity is the most immediate mechanism of adaptative response to environmental change. Studying plastic changes in response to fluctuating environments provides insights into how such adjustments may impact life history traits. Here, we used a 14-year data set of repeated body mass measurements in male eastern chipmunks (Tamias striatus) to assess the extent of plastic changes for this trait in a resource pulse ecosystem. We first determined the magnitude of variation in body mass at the population level in response to the drastic change in food resource availability from American beech tree seeds (Fagus grandifolia). Males that emerged in the spring from winter torpor following a nonmast year had a lower body mass than males emerging after a mast year, but they tended to recover this loss by mid-June. We found significant among-individual variation in spring body mass plasticity (i.e., individual-by-environment interaction). We then investigated the relationships between individual spring body mass plasticity, longevity, and lifetime reproductive success. Interestingly, heavier males lived longer than lighter males, but more plastic males had a lower longevity and lower lifetime reproductive success than less plastic males. The report of such plastic response in a stochastic resource system provides valuable insights into the interplay between the costs and benefits of phenotypic plasticity as an adaptation to environmental fluctuations.

Early-life adversity predicts performance and fitness in a wild social carnivore.

Studies on humans indicate that encountering multiple sources of adversity in childhood increases the risk of poor long-term health and premature death. Far less is known about cumulative effects of adversity during early life in wildlife. Focusing on the spotted hyena Crocuta crocuta, a social mammal with small litters, extensive maternal care, slow development and access to resources determined by social rank, we determined the contribution of ecological, maternal, social and demographic factors during early life on performance and fitness, and tested whether the impact of early-life adversity is cumulative. Using longitudinal data from 666 female hyenas in the Serengeti National Park, we determined the early growth rate, survival to adulthood, age at first reproduction (AFR), lifetime reproductive success (LRS) and longevity. We fitted multivariate models in which we tested the effects of environmental factors on these performance measures. We then constructed a cumulative adversity index and fitted models to test the effect of this index on each performance measure. Finally, the value of cumulative adversity models was tested by comparing them to multivariate and single-effect models in which the effect of each environmental factor was considered separately. High maternal rank decreased the AFR of daughters. Singleton and dominant cubs had higher growth rate than subordinate cubs, and singletons also had a higher survival chance to adulthood than subordinates. Daughters of prime age mothers had a higher growth rate, longevity and LRS. Little and heavy rainfall decreased survival to adulthood. Increasing numbers of lactating female clan members decreased growth rate, survival to adulthood and LRS. Cumulative adversity negatively affected short-term performance and LRS. Multivariate models outperformed cumulative adversity and single-effect models for all measures except for AFR and longevity, for which single-effect models performed better. Our results suggest that in some wildlife populations the combination of specific conditions in early life may matter more than the accumulation of adverse conditions as such.

Ontogenetic Expressions of Sexually Dimorphic Traits in the Skeleton Shrimp Caprella scaura (Crustacea: Amphipoda).

In sexual dimorphism, males often exhibit exaggerated characters as weapons or ornaments. Among the numerous amphipod species (Amphipoda, Crustacea) showing sexual dimorphism, caprellids (Caprellidae) are characterized by considerably larger males that possess weapons, although the developmental processes underlying these sex-related differences are largely unknown. Therefore, to clarify the process of sexual differentiation during postembryonic development in caprellids, morphometric analyses of Caprella scaura were conducted. Principal component analysis using 31 morphometric traits showed drastic allometric changes occurring at two ontogenetic body length (BL) points (i.e., 3.8 and 8.8 mm). In individuals larger than 3 mm, head spines appeared in both sexes, and penises did only in males, allowing the discrimination of juveniles from larvae. Moreover, in larger males (BL > 8.8 mm), traits used in reproductive behavior, i.e., the first antenna, second gnathopod, and first to fifth pereonites, were extremely exaggerated. Observations of pre-copulatory behavior along with morphological assays revealed that sexually mature males could be identified by the size ratio between the triangular projection and palmar spine on the propodus of the second gnathopod. In contrast, female maturation could be determined by the marginal setae of oostegites forming a brood pouch. The body size distribution of sexually mature females was concentrated within a narrow range of BLs (6-9 mm), whereas that of sexually mature males showed a broader range (BL 9-18 mm), suggesting that, in C. scaura, males continue to molt and grow even after sexual maturation via indeterminate growth, to increase their lifetime reproductive success.