Female-biased Adult Sex Ratio Research Articles

Sexual dimorphism in subterranean amphipod crustaceans covaries with subterranean habitat type.

Sexual dimorphism can evolve in response to sex-specific selection pressures that vary across habitats. We studied sexual differences in subterranean amphipods Niphargus living in shallow subterranean habitats (close to the surface), cave streams (intermediate), and cave lakes (deepest and most isolated). These three habitats differ because at greater depths there is lower food availability, reduced predation, and weaker seasonality. Additionally, species near the surface have a near-even adult sex ratio (ASR), whereas species from cave lakes have a female-biased ASR. We hypothesized (a) a decrease in sexual dimorphism from shallow subterranean habitats to cave lake species because of weaker sexual selection derived from changes in the ASR and (b) an increase in female body size in cave lakes because of stronger fecundity selection on account of oligotrophy, reduced predation, and weaker seasonality. We measured body size and two sexually dimorphic abdominal appendages for all 31 species and several behaviours related to male competition (activity, risk-taking, exploration) for 12 species. Species with an equal ASR that live close to the surface exhibited sexual dimorphism in all three morphological traits, but not in behaviour. The body size of females increased from the surface to cave lakes, but no such trend was observed in males. In cave lake species, males and females differed neither morphologically nor behaviourally. Our results are consistent with the possibility that sexual and fecundity selection covary across the three habitats, which indirectly and directly, respectively, shape the degree of sexual dimorphism in Niphargus species.

Female biased adult sex ratio in the Bronze Age cemetery of Shahr-i Sokhta (Iran) as an indicator of long distance trade and matrilocality.

This paper starts from the unusual observation of the overrepresentation of females among adults in the cemetery of Bronze Age Shahr-i Sokhta (Seistan, Iran) and explores the post marital residence pattern. By integrating taphonomical (skeletal preservation), anthropological (sex ratio [SR], sexual dimorphism, stress indicators, age at death), archeological (long distance trade indicators, habitation floor area, social role of women), and ancient DNA (heterozygosity levels in X chromosomes) data we test the hypothesis of post marital matrilocality in the site. We computed the SR (pelvis-based sex determination) in a random unpublished adult sample from the cemetery of Shahr-i Sokhta and in two samples previously published by other authors. We used comparative data on SR from: a large Supra Regional multi-chronological sample of sites, n = 47, with 8808 adult sexed individuals, from Southern Europe, Egypt, Middle East, Southern Russia; a Regional Bronze Age sample of sites (n = 10) from Bactria Margiana and Indus Valley with 1324 adult sexed individuals. We estimated the heterozygosity levels in X chromosomes compared with the rest of the autosomes on the assumption that in a matrilocal society females should show lower variability than men. Adult SR in a sample (n = 549) from Shahr-i Sokhta is 70.5, the overrepresentation of females is shared with Regional Bronze Age sites from Bactria Margiana (SR = 72.09) and Indus Valley (SR = 67.54). On the contrary, in a larger Supra Regional multi-chronological sample of sites, mean SR ranges between 112.7 (Bronze Age) and 163.1 (Middle Ages). Taphonomical and anthropological indicators do not explain the overrepresentation of female skeletons. Archeological indicators suggest a high social status of women and that the society was devoted to long range trade activities. heterozygosity levels in X chromosomes are in agreement with a matrilocal society. Indicators suggest that Bronze Age Shahr-ì Sokhta was a matrilocal society and that long distance trade was an important economic factor producing an overrepresentation of adult female skeletons in the cemetery.

A possible case of offspring sex manipulation as result of a biased adult sex ratio

Although random meiosis should prevent the facultative adjustment of offspring sex ratio, theory predicts that females should produce more of the sex with the higher reproductive value. We reported a case of offspring sex ratio manipulation in grass wrens Cistothorus platensis. Males in better body condition would have higher reproductive value than females due to the potential for social polygyny and extra-pair fertilizations. On the other hand, local demography influences reproductive strategies in grass wrens as male abundance affects both social polygyny and extra-pair paternity frequencies. We evaluated whether females bias their brood sex ratio in response to adult sex ratio and nestling body condition (a proxy for female’s prospects of producing high-quality males). Females raised more male offspring when males were less abundant in the population (female-biased adult sex ratio). However, we found no relationship between nestling body condition and brood sex ratio, suggesting that females did not bias the brood sex ratio towards males when able to raise nestlings in better body condition. Taken together, our results provide the first suggestive evidence that female birds can manipulate their offspring sex ratio in response to the adult sex ratio.

Mitonuclear Interactions and the Origin of Macaque Societies.

In most eukaryotes, aerobic respiration requires interactions between autosomally encoded genes (Ninteract genes) and mitochondrial DNA, RNA, and protein. In species where females are philopatric, contrasting distributions of genetic variation in mitochondrial and nuclear genomes create variation in mitonuclear interactions that may be subject to natural selection. To test this expectation, we turned to a group with extreme female philopatry: the macaque monkeys. We examined four genomic data sets from (1) wild caught and (2) captive populations of rhesus macaque, which is the most widely distributed nonhuman primate, and (3) the stump-tailed macaque and (4) a subspecies of longtail macaque, both of whose mitochondrial DNA is introgressed from a highly diverged ancestor. We identified atypically long runs of homozygosity, low polymorphism, high differentiation, and/or rapid protein evolution associated with Ninteract genes compared with non-Ninteract genes. These metrics suggest a subset of Ninteract genes were independently subject to atypically pervasive natural selection in multiple species. These findings suggest that natural selection on mitonuclear interactions could have influenced several aspects of macaque societies including species diversity, ecological breadth, female-biased adult sex ratio and demography, sexual dimorphism, and mitonuclear phylogenomics.

Demographic changes in an Atlantic Forest primate community following a yellow fever outbreak.

We investigated demographic changes in three primate species (Alouatta guariba, Sapajus nigritus, and Callithrix flaviceps) at the Reserva Particular do Patrimônio Natural-Feliciano Miguel Abdala,Caratinga, Minas Gerais, Brazil, following a yellow fever outbreak (YFO) by comparing their population sizes before (2015) and after the outbreak (2017-2018), and by monitoring the size, composition, and reproductive status of groups from 2017 to 2021. Comparisons of pre- and post-YFO census data indicate the A. guariba population declined by 86.6%, from an estimated minimum of 522 individuals to 70 individuals. However, by October 2021, the population had grown to at least 86 individuals, with an adult sex ratio (N = 53) that was female-biased (0.61). Eleven of the 13 groups being monitored systematically were reproductively active with high survivorship to 12 months of age. S. nigritus declined by 40%, from 377 to 226 individuals. The sex ratio of 33 adult S. nigritus is also female-biased (0.71), and at least 8 of 15 groups being monitored are reproductively active. C. flaviceps declined by 80%, from 85 individuals to the 15-17 individuals observed from 2017 to 2021. The female-biased adult sex ratio and presence of infants and juveniles in the A. guariba and S. nigritus groups are encouraging signs, but there is still great concern, especially for C. flaviceps. Continued monitoring of the demographics of these primates is needed as their persistence appears to still be at risk.

No room for males in caves: Female-biased sex ratio in subterranean amphipods of the genus Niphargus.

Sex allocation theory predicts that the proportion of daughters to sons will evolve in response to ecological conditions that determine the costs and benefits of producing each sex. All else being equal, the adult sex ratio (ASR) should also vary with ecological conditions. Many studies of subterranean species reported female-biased ASR, but no systematic study has yet been conducted. We test the hypothesis that the ASR becomes more female-biased with increased isolation from the surface. We compiled a data set of ASRs of 35species in the subterranean amphipod Niphargus, each living in one of three distinct habitats (surface-subterranean boundary, cave streams, phreatic lakes) representing an environmental gradient of increased isolation underground. The ASR was female-biased in 27 of 35species; the bias was statistically significant in 12species. We found a significant difference in the ASR among habitats after correction for phylogeny. It is most weakly female-biased at the surface-subterranean boundary and most strongly female-biased in phreatic lakes. Additional modelling suggests that the ASR has evolved towards a single value for both surface-subterranean boundary and cave stream-dwelling species, and another value for 9 of 11 phreatic lake dwellers. We suggest that a history of inbreeding in subterranean populations might lower inbreeding depression such that kin selection favours mating with siblings. This could select for a female-biased offspring sex ratio due to local mate competition among brothers. The observed patterns in sex ratios in subterranean species make them a group worthy of more attention from those interested in sex allocation theory.

Home range use in the West Australian seahorse Hippocampus subelongatus is influenced by sex and partner\u2019s home range but not by body size or paired status

Genetic monogamy is the rule for many species of seahorse, including the West Australian seahorse Hippocampus subelongatus. In this paper, we revisit mark-recapture and genetic data of H. subelongatus, allowing a detailed characterization of movement distances, home range sizes and home range overlaps for each individual of known sex, paired status (paired or unpaired) and body size. As predicted, we find that females have larger home ranges and move greater distances compared to males. We also confirm our prediction that the home ranges of pair-bonded individuals (members of a pair known to reproduce together) overlap more on average than home ranges of randomly chosen individuals of the opposite or same sex. Both sexes, regardless of paired status, had home ranges that overlapped with, on average, 6–10 opposite-sex individuals. The average overlap area among female home ranges was significantly larger than the overlap among male home ranges, probably reflecting females having larger home ranges combined with a female biased adult sex ratio. Despite a prediction that unpaired individuals would need to move around to find a mate, we find no evidence that unpaired members of either sex moved more than paired individuals of the same sex. We also find no effect of body size on home range size, distance moved or number of other individuals with which a home range overlapped. These patterns of movement and overlap in home ranges among individuals of both sexes suggest that low mate availability is not a likely explanation for the maintenance of monogamy in the West Australian seahorse.

Female-Biased Adult Sex Ratio in Relation to Sex-Specific Adult and First-Year Survival, Fledgling Sex Ratio and Dispersal in a Migratory Passerine

The relative number of adult males to females in a population, the adult sex ratio (ASR), is an important demographic variable in populations, which plays a role in, for instance, mating systems and viability of populations. The origins of variation in ASR between populations are often unclear. A skewed ASR in a local population may result from a skewed sex ratio among fledglings, sex differences in survival or dispersal or a combination of these. Meta-analyses show that ASRs across bird species are mostly male-biased and that male-biased survival is the main contributor to this bias. Here, we examine the origins of a female-biased ASR in a small population (3–24 breeding females in 2007–2017) of a passerine, the Northern Wheatear Oenanthe oenanthe. Because 95% of all fledglings were sexed, 88% of the individuals were colour-banded and because this species is highly philopatric, we could accurately examine: (1) fledgling sex ratio, (2) sex-specific first year and adult apparent survival and (3) sex-specific dispersal. It appeared that survival was not sex-biased for either age cohort but that fledgling sex ratio was male-biased, especially so among late broods. In addition, natal dispersal out of the breeding population was likely male-biased, in contrast to the general pattern of female-bias in avian dispersal. So, perhaps females are more common in our relict population solely because more males disperse from the population than females, without being compensated by male-biased fledgling sex ratio or survival and without compensation from immigrating males. Likely, the population sex ratio (including both adults and young) at the end of the breeding season is male-biased, because of a pronounced male-biased sex ratio among fledglings from late nests. Perhaps this male-bias induces prospecting young males to settle elsewhere to avoid competition next year, but at the same time convinces prospecting females to settle particularly in this area.

Causes and consequences of an unusually male-biased adult sex ratio in an unmanaged feral horse population.

The adult sex ratio (ASR) is important within ecology due to its predicted effects on behaviour, demography and evolution, but research examining the causes and consequences of ASR bias have lagged behind the studies of sex ratios at earlier life stages. Although ungulate ASR is relatively well-studied, exceptions to the usual female-biased ASR challenge our understanding of the underlying drivers of biased ASR and provide an opportunity to better understand its consequences. Some feral ungulate populations, including multiple horse populations, exhibit unusually male-biased ASR. For example, research suggests that the feral horse Equus ferus caballus population on Sable Island, Nova Scotia, Canada may exhibit a male-biased ASR. Such exceptions to the rule provide a valuable opportunity to reveal the contributions of environmental context and trait differences to ASR bias. We aimed to test for bias in Sable Island horse ASR, identify the demographic drivers of bias, and explore its demographic and social consequences. To do this, we used life history, movement and group membership information for hundreds of horses followed through a long-term individual-based study between 2007 and 2018. Sable Island horse ASR is male biased and this skew has increased over time, reaching 62% male in 2018. Our life table response experiment suggested that ASR skew was driven predominantly by male-biased adult survival. Further analyses pointed to sex-biased survival being driven by reduced female survival post-reproduction. Male-biased ASR was associated with reduced harem sizes, an increase in the number of social groups on the island, and reduced reproduction in young females. Our results support the idea that male-biased ASR in feral ungulate populations may be caused by a combination of high population density and high reproductive output. We suggest that female-biased mortality may be caused by females continuing to reproduce at high density, and thus being more susceptible to resource shortages. Thus, our results highlight the strong context dependence of ASR. Furthermore, our work indicates the potential for ASR to substantially alter a population's social organisation. Such changes in social structure could have knock-on consequences for demography by altering the formation/stability of social relationships, or competition for matings.

Trade-off between mate choice speed and decision accuracy under mating competition in female sand gobies

Variation in female mating performance can affect the direction and rate of evolution through sexual selection. The social environment determines the availability of mates and the competitive situation, and hence can influence mating strategies. However, these effects have to be considered within time constraints. Here we show that female sand gobies exposed to same-sex competitors (female-biased adult sex ratio, ASR) for a week before having physical access to males (i.e., a delayed male access; referred as prior exposure treatment) were more actively associated with the preferred male and took faster spawning decisions. However, these females mated more frequently with males with traits that did not ensure high egg survival. On the other extreme, females exposed to low same sex competition (male-biased ASR) simultaneously to the mate choice (i.e., an immediate access to males; referred as simultaneous exposure treatment) took more time to make their spawning decision. They also associated and spawned more often with males with good parental skills (i.e., males exhibiting more egg fanning behaviour). These male traits are associated with higher survival and better development of eggs. Our results provide experimental support for a trade-off between mate choice accuracy and speed, which depended on mating competition.

Sex Ratio of Small Hive Beetles: The Role of Pupation and Adult Longevity.

The sex ratio of sexually reproducing animal species tends to be 1:1, which is known as Fisher’s principle. However, differential mortality and intraspecific competition during pupation can result in a biased adult sex ratio in insects. The female-biased sex ratio of small hive beetles (SHBs) is known from both laboratory and field studies, but the underlying reasons are not well understood. Here, we used laboratory mass and individual pupation to test if differential mortality between sexes and/or intraspecific interactions can explain this sex ratio. The data show a significant female-biased adult sex ratio in both mass and individual rearing, even when assuming that all dead individuals were males. Our results therefore suggest that neither differential mortality during pupation nor intraspecific interactions are likely to explain the female-biased sex ratio of freshly emerged adult SHBs. We regard it as more likely that either competition during the larval feeding stage or genetic mechanisms are involved. In addition, we compared our data with previously published data on the sex ratio of both freshly emerged and field-collected SHBs to investigate possible gender differences in adult longevity. The data show a significantly greater female bias in the sex ratio upon emergence, compared to field-collected SHBs, suggesting that adult females have a shorter longevity.

Adult sex-ratio in ostracods and its implications for sexual selection

ABSTRACTThe Ostracoda – ubiquitous aquatic micro-crustaceans – show an exceptionally high incidence of female-biased adult sex ratio. Intraspecific sex ratio is known to vary in extant species and yet in the fossil record a species’ adult sex ratio can be highly stable across time. Sex ratio conditions the intensity of sexual selection and influences which sex undergoes stronger selective pressure. However, the impact of variation in spatial and temporal intraspecific sex ratio on the evolution of sexual selection remains an open question, calling for further investigations on the factors controlling adaptive sex ratio. This mini-review aims to introduce the system, and explores some of the key literature addressing factors influencing intraspecific variation in adult sex ratio (ASR) and its implication in the intensity of sexual selection and evolution of mating systems.

Prolonged milk provisioning in a jumping spider.

Lactation is a mammalian attribute, and the few known nonmammal examples have distinctly different modalities. We document here milk provisioning in a jumping spider, which compares functionally and behaviorally to lactation in mammals. The spiderlings ingest nutritious milk droplets secreted from the mother's epigastric furrow until the subadult stage. Milk is indispensable for offspring survival in the early stages and complements their foraging in later stages. Maternal care, as for some long-lived vertebrates, continues after the offspring reach maturity. Furthermore, a female-biased adult sex ratio is acquired only when the mother is present. These findings demonstrate that mammal-like milk provisioning and parental care for sexually mature offspring have also evolved in invertebrates, encouraging a reevaluation of their occurrence across the animal kingdom, especially in invertebrates.

Social determinants of adult sex ratios and racial/ethnic disparities in transmission of HIV and other sexually transmitted infections in the USA.

In Black population centres in the USA, adult sex ratios (ASRs) are strongly female-biased primarily due to high male incarceration and early mortality rates. I explore the system of social determinants that shape these ASRs, and describe their apparent consequences. Evidence suggests that female-biased ASRs play a role, along with racial residential segregation, to increase mixing between core and peripheral members of sexual networks, facilitating transmission of human immunodeficiency virus and other sexually transmitted infections. Unique historical factors underlie Black male incarceration and mortality rates in the USA, making comparisons with other groups or other countries challenging.This article is part of the themed issue 'Adult sex ratios and reproductive decisions: a critical re-examination of sex differences in human and animal societies'.

Combining genetic structure and demographic analyses to estimate persistence in endangered Key deer (Odocoileus virginianus clavium)

Recent improvements in genetic analyses have paved the way in using molecular data to answer questions regarding evolutionary history, genetic structure, and demography. Key deer are a federally endangered subspecies assumed to be genetically unique, homogeneous, and have a female-biased population of approximately 900 deer. We used 985 bp of the mitochondrial cytochrome b gene and 12 microsatellite loci to test two hypotheses: (1) that Key deer are isolated and have reduced diversity compared to mainland deer and (2) that isolation of the Florida Keys has led to a small population size and a high risk of extinction. Our results indicate that Key deer are indeed genetically isolated from mainland white-tailed deer and that there is a lack of genetic substructure between islands. While Key deer exhibit reduced levels of genetic diversity compared to their mainland counterparts, they contain enough diversity to uniquely identify individual deer. Based on genetic identification, we estimated a census size of around 1000 individuals with a heavily skewed female-biased adult sex ratio. Furthermore, our genetic and contemporary demographic data were used to generate a species persistence model of the Key deer. Sensitivity tests within the population viability analysis brought to light the importance of fetal sex ratio and female survival as the primary factors at risk of driving the subspecies to extinction. This study serves as a prime example of how persistence models can be used to evaluate population viability in natural populations of endangered organisms.

Crowd control: sex ratio affects sexually selected cuticular hydrocarbons in male Drosophila serrata.

Although it is advantageous for males to express costly sexually selected signals when females are present, they may also benefit from suppressing these signals to avoid costly interactions with rival males. Cuticular chemical profiles frequently function as insect sexual signals; however, few studies have asked whether males alter these signals in response to their social environment. In Drosophila serrata, an Australian fly, there is sexual selection for a multivariate combination of male cuticular hydrocarbons (CHCs). Here, we show that the ratio of females to males that an adult male experiences has a strong effect on his CHC expression, with female-biased adult sex ratios eliciting greater expression of CHC profiles associated with higher male mating success. Classical models predict that male reproductive investment should be highest when there is a small but nonzero number of rivals, but we found that males expressed the most attractive combination of CHCs when there were no rivals. We found that male CHCs were highly sensitive to adult sex ratio, with males expressing higher values of CHC profiles associated with greater mating success as the ratio of females to males increased. Moreover, sex ratio has a stronger effect on male CHC expression than adult density. Finally, we explore whether sex ratio affects the variance among a group of males in their CHC expression, as might be expected if individuals respond differently to a given social environment, but find little effect. Our results reveal that subtle differences in social environment can induce plasticity in male chemical signal expression.

Female-biased sex ratio, polygyny, and persistence in the endangered Southwestern Willow Flycatcher (Empidonax traillii extimus)

ABSTRACT Demographic changes in populations, such as skewed sex ratios, are of concern to conservationists, especially in small populations in which stochastic and other events can produce declines leading to extirpation. We documented a decline in one of the few remaining populations of Southwestern Willow Flycatcher (Empidonax traillii extimus) in southern California, USA, which dropped from 40 to 5 adults between 2000 and 2015. Declines were unequal between sexes (94% for males, 82% for females). Adult sex ratios were female-biased in 10 of 16 yr. The proportion of paired males that were polygynous ranged from 0% to 100%, depending on the ratio of females to males in the adult population. Some males paired with up to 5 females simultaneously. We investigated the role of nestling sex ratio in the female-biased adult sex ratio by using genetic techniques to determine sex from blood samples collected from 162 nestlings in 72 nests from 2002 to 2009. Both population-level and within-brood nestling sex rati...

Status of Golden Jackal Canis aureus and ungulates in a small enclosed area- Van Vihar National Park, Madhya Pradesh, India

We estimated densities of Golden Jackals and five ungulate species in Van Vihar National Park, Madhya Pradesh, India. It is an enclosed area of about 4.45km², out of which 3.5km² is available for free ranging animals. Twenty-six transects with a combined length of 22.6km and an effort of 50.2km were walked. A total of 1079 animal detections belonging to six different species were made. The density of jackals was (17±3.8SE)/km 2 . Among the ungulates, chital had the highest density (118±18.8SE)/km 2 followed by Sambar (34.1±6.9 SE)/km 2 , Nilgai (13.1±2.8SE)/km 2 , Blackbuck (6.6±1.5 SE)/km 2 and Wild Pig (3.7±0.8 SE)/km 2 . The ungulate biomass was found to be (12979.2±2463.26 SE)kg/km 2 . Chital biomass was the highest at (5574.2±886.58 SE)kg/km 2 , followed by Sambar biomass of (4569.4±913.75 SE)kg/km 2 , Nilgai (2358±523.24 SE)kg/km 2 , Blackbuck (211.2±66.18 SE)kg/km 2 and Wild Pig (118.4±28.37 SE)kg/km 2 . The sex ratio was calculated and most ungulates had female-biased adult sex ratio.

Male-directed infanticide in spider monkeys (Ateles spp.).

Infanticide is considered a conspicuous expression of sexual conflict amongst mammals, including at least 35 primate species. Here we describe two suspected and one attempted case of intragroup infanticide in spider monkeys that augment five prior cases of observed or suspected infanticide in this genus. Contrary to the typical pattern of infanticide seen in most primate societies, where infants are killed by conspecifics independent of their sex, all eight cases of observed or suspected infanticide in spider monkeys have been directed toward male infants within their first weeks of life. Moreover, although data are still scant, infanticides seem to be perpetrated exclusively by adult males against infants from their own social groups and are not associated with male takeovers or a sudden rise in male dominance rank. Although the slow reproductive cycles of spider monkeys might favor the presence of infanticide because of the potential to shorten females' interbirth intervals, infanticide is nonetheless uncommon among spider monkeys, and patterns of male-directed infanticide are not yet understood. We suggest that given the potentially close genetic relationships among adult males within spider monkey groups, and the need for males to cooperate with one another in territorial interactions with other groups of related males, infanticide may be expected to occur primarily where the level of intragroup competition among males outweighs that of competition between social groups. Finally, we suggest that infanticide in spider monkeys may be more prevalent than previously thought, given that it may be difficult for observers to witness cases of infanticide or suspected infanticide that occur soon after birth in taxa that are characterized by high levels of fission-fusion dynamics. Early, undetected, male-biased infanticide could influence the composition of spider monkey groups and contribute to the female-biased adult sex ratios often reported for this genus.

Reduced representation genome sequencing suggests low diversity on the sex chromosomes of tonkean macaque monkeys.

In species with separate sexes, social systems can differ in the relative variances of male versus female reproductive success. Papionin monkeys (macaques, mangabeys, mandrills, drills, baboons, and geladas) exhibit hallmarks of a high variance in male reproductive success, including a female-biased adult sex ratio and prominent sexual dimorphism. To explore the potential genomic consequences of such sex differences, we used a reduced representation genome sequencing approach to quantifying polymorphism at sites on autosomes and sex chromosomes of the tonkean macaque (Macaca tonkeana), a species endemic to the Indonesian island of Sulawesi. The ratio of nucleotide diversity of the X chromosome to that of the autosomes was less than the value (0.75) expected with a 1:1 sex ratio and no sex differences in the variance in reproductive success. However, the significance of this difference was dependent on which outgroup was used to standardize diversity levels. Using a new model that includes the effects of varying population size, sex differences in mutation rate between the autosomes and X chromosome, and GC-biased gene conversion (gBGC) or selection on GC content, we found that the maximum-likelihood estimate of the ratio of effective population size of the X chromosome to that of the autosomes was 0.68, which did not differ significantly from 0.75. We also found evidence for 1) a higher level of purifying selection on genic than nongenic regions, 2) gBGC or natural selection favoring increased GC content, 3) a dynamic demography characterized by population growth and contraction, 4) a higher mutation rate in males than females, and 5) a very low polymorphism level on the Y chromosome. These findings shed light on the population genomic consequences of sex differences in the variance in reproductive success, which appear to be modest in the tonkean macaque; they also suggest the occurrence of hitchhiking on the Y chromosome.