Male-biased Sex Ratio Research Articles

Heavier females produce more sons in a low‐density population of red deer

According to the Trivers-Willard hypothesis, females of polygynous mammal species can adaptively modify offspring sex ratio, which would provide them evolutionary benefits in terms of a higher number of grand-offspring. In red deer (Cervus elaphus), a species where the reproductive success of males is significantly more variable than that of females, mothers that are in good condition are expected to have a male-biased offspring sex ratio but only in a population below carrying capacity. We aimed at testing the Trivers-Willard hypothesis on a low-density population of red deer in large woodlands of north-eastern Poland. The overall foetal sex ratio did not differ from parity. The mother's probability of having male offspring increased with her body mass independently of age and population density. Such condition-based adjustment of the offspring sex ratio in female red deer from a low-density population conforms to the Trivers–Willard hypothesis. However, full support of Trivers–Willard predictions would require further investigations comparing survival and reproductive output of male and female offspring.

The male handicap: male-biased mortality explains skewed sex ratios in brown trout embryos.

Juvenile sex ratios are often assumed to be equal for many species with genetic sex determination, but this has rarely been tested in fish embryos due to their small size and absence of sex-specific markers. We artificially crossed three populations of brown trout and used a recently developed genetic marker for sexing the offspring of both pure and hybrid crosses. Sex ratios (SR = proportion of males) varied widely one month after hatching ranging from 0.15 to 0.90 (mean = 0.39 ± 0.03). Families with high survival tended to produce balanced or male-biased sex ratios, but SR was significantly female-biased when survival was low, suggesting that males sustain higher mortality during development. No difference in SR was found between pure and hybrid families, but the existence of sire × dam interactions suggests that genetic incompatibility may play a role in determining sex ratios. Our findings have implications for animal breeding and conservation because skewed sex ratios will tend to reduce effective population size and bias selection estimates.

Sex Ratio Elasticity Influences the Selection of Sex Ratio Strategy.

There are three sex ratio strategies (SRS) in nature—male-biased sex ratio, female-biased sex ratio and, equal sex ratio. It was R. A. Fisher who first explained why most species in nature display a sex ratio of ½. Consequent SRS theories such as Hamilton’s local mate competition (LMC) and Clark’s local resource competition (LRC) separately explained the observed deviations from the seemingly universal 1:1 ratio. However, to the best of our knowledge, there is not yet a unified theory that accounts for the mechanisms of the three SRS. Here, we introduce the price elasticity theory in economics to define sex ratio elasticity (SRE), and present an analytical model that derives three SRSs based on the following assumption: simultaneously existing competitions for both resources A and resources B influence the level of SRE in both sexes differently. Consequently, it is the difference (between two sexes) in the level of their sex ratio elasticity that leads to three different SRS. Our analytical results demonstrate that the elasticity-based model not only reveals a highly plausible mechanism that explains the evolution of SRS in nature, but also offers a novel framework for unifying two major classical theories (i.e., LMC & LRC) in the field of SRS research.

Virility does not imply immensity: Testis size, accessory gland size and ejaculate depletion pattern do not evolve in response to experimental manipulation of sex ratio in Drosophila melanogaster

Sperm competition theory predicts that with increase in sperm competition, males either invest more in reproductive organ(s) and/or improve ejaculate investment. We test this idea using experimental evolution in Drosophila melanogaster. We maintained replicate populations of Drosophila melanogaster under male (M) and female (F) biased sex ratio regimes for more than a hundred generations with the result that males from the M regime evolved higher sperm competitive abilities relative to males from the F regime. In the present study, we measured the testes and the accessory gland size of virgin and singly mated males from the M and F regimes. The M and F males do not differ in either testis or accessory gland size. Additionally, ejaculate investment is not different in the M and F males, as measured by reduction in testis and accessory gland sizes. Thus, contrary to theoretical prediction and evidence from other species, we found that evolved differences in sperm competitive ability are not necessarily due to evolution of testis/accessory gland size or strategic ejaculate investment in these populations.

Ecology and Demography of Free-Roaming Domestic Dogs in Rural Villages near Serengeti National Park in Tanzania.

Free-roaming dogs (Canis lupus familiaris) are of public health and conservation concern because of their potential to transmit diseases, such as rabies, to both people and wildlife. Understanding domestic dog population dynamics and how they could potentially be impacted by interventions, such as rabies vaccination, is vital for such disease control efforts. For four years, we measured demographic data on 2,649 free-roaming domestic dogs in four rural villages in Tanzania: two villages with and two without a rabies vaccination campaign. We examined the effects of body condition, sex, age and village on survivorship and reproduction. Furthermore, we compared sources of mortality among villages. We found that adult dogs (>12mos) had higher survival than puppies in all villages. We observed a male-biased sex ratio across all age classes. Overall survival in one non-vaccination village was lower than in the other three villages, all of which had similar survival probabilities. In all villages, dogs in poor body condition had lower survival than dogs in ideal body condition. Sickness and spotted hyena (Crocuta crocuta) predation were the two main causes of dog death. Within vaccination villages, vaccinated dogs had higher survivorship than unvaccinated dogs. Dog population growth, however, was similar in all the villages suggesting village characteristics and ownership practices likely have a greater impact on overall dog population dynamics than vaccination. Free-roaming domestic dogs in rural communities exist in the context of their human owners as well as the surrounding wildlife. Our results did not reveal a clear effect of vaccination programs on domestic dog population dynamics. An investigation of the role of dogs and their care within these communities could provide additional insight for planning and implementing rabies control measures such as mass dog vaccination.

Male-biased operational sex ratios and the Viking phenomenon: an evolutionary anthropological perspective on Late Iron Age Scandinavian raiding

In this paper, we use a combination of evolutionary theory, ethnographic data, written sources, and archaeological evidence to develop a new explanation for the origins of Viking raiding. Our argum ...

Androgenesis: where males hijack eggs to clone themselves.

Androgenesis is a form of quasi-sexual reproduction in which a male is the sole source of the nuclear genetic material in the embryo. Two types of androgenesis occur in nature. Under the first type, females produce eggs without a nucleus and the embryo develops from the male gamete following fertilization. Evolution of this type of androgenesis is poorly understood as the parent responsible for androgenesis (the mother) gains no benefit from it. Ultimate factors driving the evolution of the second type of androgenesis are better understood. In this case, a zygote is formed between a male and a female gamete, but the female genome is eliminated. When rare, androgenesis with genome elimination is favoured because an androgenesis-determining allele has twice the reproductive success of an allele that determines sexual reproduction. Paradoxically, except in hermaphrodites, a successful androgenetic strain can drive such a male-biased sex ratio that the population goes extinct. This likely explains why androgenesis with genome elimination appears to be rarer than androgenesis via non-nucleate eggs, although both forms are either very rare or remain largely undetected in nature. Nonetheless, some highly invasive species including ants and freshwater clams are androgenetic, for reasons that are largely unexplained.This article is part of the themed issue 'Weird sex: the underappreciated diversity of sexual reproduction'.

Genetic assessment of captive red panda (Ailurus fulgens) population.

Red panda (Ailurus fulgens) is threatened across its range by detrimental human activities and rapid habitat changes necessitating captive breeding programs in various zoos globally to save this flagship species from extinction. One of the ultimate aims of ex situ conservation is reintroduction of endangered animals into their natural habitats while maintaining 90 % of the founder genetic diversity. Advances in molecular genetics and microsatellite genotyping techniques make it possible to accurately estimate genetic diversity of captive animals of unknown ancestry. Here we assess genetic diversity of the red panda population in Padmaja Naidu Himalayan Zoological Park, Darjeeling, which plays a pivotal role in ex situ conservation of red panda in India. We generated microsatellite genotypes of fifteen red pandas with a set of fourteen loci. This population is genetically diverse with 68 % observed heterozygosity (HO) and mean inbreeding (FIS) coefficient of 0.05. However population viability analysis reveals that this population has a very low survival probability (<2 %) and will rapidly loose its genetic diversity to 37 % mainly due to small population size and skewed male-biased sex ratio. Regular supplementation with a pair of adult individuals every five years will increase survival probability and genetic diversity to 99 and 61 % respectively and will also support future harvesting of individuals for reintroduction into the wild and exchange with other zoos.

Local resource competition affects sex allocation in a bird: experimental evidence

Sex allocation theory as applied to local resource competition (LRC) predicts that parents should skew investment towards philopatric offspring when local resources are abundant. Alternatively, parents should allocate resources to the dispersing sex when local resources are deficient in order to limit competition among relatives. Nest sites have been shown to be the primary factor limiting populations of secondary cavity-nesting birds. In this study, we manipulated nestbox density to test its effect on the sex allocation patterns of the great tit, Parus major, a species in which female offspring are more likely to disperse than male offspring. We also investigated the relationship between the brood sex ratio and the time of breeding, which has been shown in many studies to influence sex allocation. Consistent with the LRC prediction, parents invested more in male offspring and produced a male-biased sex ratio in the area where nestboxes were abundant. In our study, the reproductive success of great tits declined as the season progressed; however, the time of breeding had no effect on the sex ratio of the offspring. Overall, the results of our study suggest that nestbox availability can influence sex allocation in great tits, and may also represent a relatively common phenomenon in other secondary cavity-nesting birds.

Demographic Analysis of Sex Ratio on Population Growth of Bactrocera dorsalis (Diptera: Tephritidae) With Discussion of Control Efficacy Using Male Annihilation.

The life table data for the oriental fruit fly, Bactrocera dorsalis (Hendel), at different adult sex ratios (1♀: 1♂, 1♀: 50♂, 50♀: 1♂ free-choice mating, and 50♀: 1♂ no-choice mating) were collected to determine the effects of sex-ratio manipulation on current pest control procedures. At 1♀: 1♂, females mated, on average, 2.3 times during their lifetime with a mean fecundity (F) of 1,122 eggs. The net reproductive rate (R0), intrinsic rate of increase (r), finite rate (λ), and mean generation time (T) were 561.0 offspring, 0.1693 d- 1, 1.1844 d- 1, and 37.4 d, respectively. At 50♀: 1♂ free-choice mating, males mated 46.7 times during their lifetime, while at 50♀: 1♂ no-choice mating, males mated on average 50 times during their lifetime, and all females mating only once in both treatments. The values for F, r, and λ were significantly lower for both 50♀: 1♂ treatments than those in the 1♀: 1♂ group; the R0 values, however, were either equal to or even higher than those in the 1♀: 1♂ treatment. In the male-biased sex ratio (1♀: 50♂), fecundity was the highest (1,610 eggs) and female average life span the longest (166 d), while the R0 was the lowest (31.6 offspring) among all treatments. Population projections showed that even at a sex ratio of 50♀: 1♂, B. dorsalis could still produce a large number of offspring. These findings demonstrate that management strategies for controlling B. dorsalis could be properly evaluated by using demographic methods. Because female annihilation appears to be a more effective control strategy, it should be considered as a viable alternative.

Marriage Markets and Male Mating Effort: Violence and Crime Are Elevated Where Men Are Rare.

The negative social outcomes in populations with male-biased sex ratios are a growing concern. In general, the expectation is of heightened violence as a result of excess men engaging in antisocial behavior and crime, thereby threatening societal stability. While intuitive, these claims are largely unsupported in the literature. Using mating market theory as our guide, we examine indicators of male mating effort, including (1) violent competition between men (homicide, aggravated assault) and (2) indicators of uncommitted sexual behavior (rape, sex offenses, and prostitution). Our unit of analysis is U.S. county-level data. We find that counties with more men have lower rates of crime and violent behavior. Our findings challenge conventional claims of male excess leading to elevated levels of violence. Instead, in support of mating market predictions, we find that criminal and violent behavior related to male mating effort is least common in male-biased sex ratios. We discuss the implications of our findings for public policy regarding incarceration and criminal behavior.

Competitive males have higher quality sperm in a monogamous social bee.

BackgroundReproductive success is determined by the interplay between mating and fertilization success. In social insect species with male-biased sex ratios and queen monogamy, males face particularly strong pre-copulatory sexual selection since they must compete with thousands of other males for a unique mating opportunity. Ejaculate quality is also expected to be under selection, because queens are long-lived and store sperm for life, so males with higher quality ejaculates are expected to provide queens with larger and longer-lived colonies, which in turn may produce more daughter queens (the only direct fitness gains of haplodiploid males). Considering the action of pre and post-copulatory sexual selection on male traits, three scenarios might thus be expected: positive, negative or no association between male mating ability and fertilization success. Here we explored these scenarios in the stingless bee Scaptotrigona aff. depilis, where males gather in large aggregations and queens mate with a single male. Male mating ability was assessed through the capacity of a male to reach an aggregation and persist on it; while sperm viability, sperm number, and sperm morphology were used as proxies for sperm quality.ResultsSperm viability was associated with persistence time in the aggregation, and males that persisted longer presented shorter spermatozoa and higher variation in sperm length than recently arrived males. However, sperm traits of males that reached aggregations did not differ from those of males collected inside their colonies. In addition, males that persisted longer in aggregations were smaller than other males. Male size and sperm viability were not correlated, suggesting that the observed patterns were not due to trade-offs in male resource allocation.ConclusionsPersistence in male aggregations thus seems to select more competitive males with higher quality sperm. Our work is the first one to reveal an association between male competitive ability and fertilization success in a monogamous social insect. This finding sheds important light on the evolution of male traits in social insects and the general mechanisms of sexual selection.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0765-2) contains supplementary material, which is available to authorized users.

Chastity Beliefs, Female Neglect, Academic Achievement, and Psychological Well-Being of Adolescent Girls in Tamilnadu, India

In this study, we examined the relationship between chastity beliefs and psychological well-being among adolescent girls with a specific focus on culturally valued beliefs about gender in communities with a history of extreme female neglect in Tamilnadu, India. We predicted that endorsing prescriptive gender norms would be positively related to resilience and lower levels of depressive symptoms and academic achievement. We found that, for adolescent girls (N = 250), chastity beliefs were positively related to resilience (b =.18, p <.01) and negatively related to depression (b = -.24, p <.001) and academic performance (b = -.40, p <.001). The ecological context of male-biased sex ratios shapes the lives of adolescent girls in complex ways. Although endorsing culturally valued gender beliefs enhanced the well-being of adolescent girls, it was also negatively correlated to their academic achievement. The cultural psychological implications of our findings for reproductive health education in India are discussed.

Patterns of Family Formation in Response to Sex Ratio Variation.

The impact that unbalanced sex ratios have on health and societal outcomes is of mounting contemporary concern. However, it is increasingly unclear whether it is male- or female-biased sex ratios that are associated with family and social instability. From a socio-demographic perspective, male-biased sex ratios leave many men unable to find a mate, elevating competition among males, disrupting family formation and negatively affecting social stability. In contrast, from a mating-market perspective, males are expected to be less willing to marry and commit to a family when the sex ratio is female-biased and males are rare. Here we use U.S. data to evaluate predictions from these competing frameworks by testing the relationship between the adult sex ratio and measures of family formation. We find that when women are rare men are more likely to marry, be part of a family and be sexually committed to a single partner. Our results do not support claims that male-biased sex ratios lead to negative family outcomes due to a surplus of unmarried men. Rather, our results highlight the need to pay increased attention to female-biased sex ratios.

Comparative chronic toxicity of imidacloprid, clothianidin, and thiamethoxam to Chironomus dilutus and estimation of toxic equivalency factors.

Nontarget aquatic insects are susceptible to chronic neonicotinoid insecticide exposure during the early stages of development from repeated runoff events and prolonged persistence of these chemicals. Investigations on the chronic toxicity of neonicotinoids to aquatic invertebrates have been limited to a few species and under different laboratory conditions that often preclude direct comparisons of the relative toxicity of different compounds. In the present study, full life-cycle toxicity tests using Chironomus dilutus were performed to compare the toxicity of 3 commonly used neonicotinoids: imidacloprid, clothianidin, and thiamethoxam. Test conditions followed a static-renewal exposure protocol in which lethal and sublethal endpoints were assessed on days 14 and 40. Reduced emergence success, advanced emergence timing, and male-biased sex ratios were sensitive responses to low-level neonicotinoid exposure. The 14-d median lethal concentrations for imidacloprid, clothianidin, and thiamethoxam were 1.52 μg/L, 2.41 μg/L, and 23.60 μg/L, respectively. The 40-d median effect concentrations (emergence) for imidacloprid, clothianidin, and thiamethoxam were 0.39 μg/L, 0.28 μg/L, and 4.13 μg/L, respectively. Toxic equivalence relative to imidacloprid was estimated through a 3-point response average of equivalencies calculated at 20%, 50%, and 90% lethal and effect concentrations. Relative to imidacloprid (toxic equivalency factor [TEF] = 1.0), chronic (lethality) 14-d TEFs for clothianidin and thiamethoxam were 1.05 and 0.14, respectively, and chronic (emergence inhibition) 40-d TEFs were 1.62 and 0.11, respectively. These population-relevant endpoints and TEFs suggest that imidacloprid and clothianidin exert comparable chronic toxicity to C. dilutus, whereas thiamethoxam induced comparable effects only at concentrations an order of magnitude higher. However, the authors caution that under field conditions, thiamethoxam readily degrades to clothianidin, thereby likely enhancing toxicity. Environ Toxicol Chem 2017;36:372-382. © 2016 SETAC.

Evolutionary implications for the determination of gametocyte sex ratios under fecundity variation for the malaria parasite

We investigate sex ratio determination strategies for the Malaria parasite based on putative changes in its male fecundity over the lifetime of an infection, and how such strategies might have evolved. We model fitness using the incomplete fertilization limit developed in Teboh-Ewungkem and Yuster (2010). We divide the infection lifetime of a strain into two periods, assume each human is infected by two different strains, and assume that there are two different strategies present among the many strains in the general malaria parasite population. A unique parameter dependent ESS exists for all parameter values in both of our main models, with many such strategies unbeatable. These strategies produce both male and female biased population sex ratios with female bias predominating over most of the parameter space. The first model (SKM) suggests that strains without the ability to detect characteristics of other strains present could still have evolved strategies to vary sex ratio over their lifetimes, and the second model (DKM) suggests strains with detection abilities might have evolved after that. Our analysis suggests that once the ability to detect the population sizes and fecundities of other strains has developed, detection of their sex ratio choices confers no additional selective advantage in that a DKM ESS is still an ESS among sex ratio detecting strategies. The sex ratio choices for each DKM ESS are given by the equilibrium values of the parameter equivalent sex ratio detecting strategy described in Teboh-Ewungkem and Wang (2012), in the case where two strains employing that strategy encounter each other.

Climate change perils for dioecious plant species.

Climate change, particularly increased aridity, poses a significant threat to plants and the biotic communities they support. Dioecious species may be especially vulnerable to climate change given that they often exhibit spatial segregation of the sexes, reinforced by physiological and morphological specialization of each sex to different microhabitats. In dimorphic species, the overexpression of a trait by one gender versus the other may become suppressed in future climates. Data suggest that males will generally be less sensitive to increased aridity than co-occurring females and, consequently, extreme male-biased sex ratios are possible in a significant number of populations. The effects of male-biased sex ratios are likely to cascade to dependent community members, especially those that are specialized on one sex.

Contrasting responses in the growth and energy utilization properties of sympatric Populus and Salix to different altitudes: implications for sexual dimorphism in Salicaceae.

An interesting ecological and evolutionary puzzle arises from the observations of male-biased sex ratios in genus Populus, whereas in the taxonomically related Salix, females are generally more dominant. In the present study, we combined results from a field investigation into the sex ratios of the Salicaceous species along an altitudinal gradient on Gongga Mountain, and a pot experiment by monitoring growth and energy utilization properties to elucidate the mechanisms governing sexual dimorphism. At middle altitudes 2000 and 2300 m, the sex ratios were consistent with a 1:1 equilibrium in sympatric Populus purdomii and Salix magnifica. However, at the lower and higher ends of the altitudinal gradient, skewed sex ratios were observed. For example, the male:female ratios were 1.33 and 2.36 in P. purdomii at 1700 and 2600 m respectively; for S. magnifica the ratio was 0.62 at 2600 m. At 2300 m, the pot-grown seedlings of both species exhibited the highest biomass accumulation and total leaf area, simultaneously with the balanced sex ratios in the field. At 3300 m, the specific leaf area in male P. purdomii was 23.9% higher than that of females, which may be the morphological cause for the observed 19.3% higher nitrogen allocation to Rubisco, and 20.6% lower allocation to cell walls. As such, male P. purdomii showed a 32.9% higher foliar photosynthetic capacity, concomitant with a 12.0% lower construction cost. These properties resulted in higher photosynthetic nitrogen- and energy-use efficiencies, and shorter payback time (24.4 vs 40.1 days), the time span that a leaf must photosynthesize to amortize the carbon investment. Our results thus suggested that male P. purdomii evolved a quicker energy-return strategy. Consequently, these superior energy gain-cost related traits and the higher total leaf area contributed to the higher growth rate and tolerance in stress-prone environments, which might, in part, shed new light on the male-biased sex ratios in Populus. However, no significant sexual difference was observed in S. magnifica for all the above parameters, thereby implying that the female-biased sex ratios in Salix cannot be explained in terms of the energy-use properties studied here.

Causes and consequences of spatial variation in sex ratios in a declining bird species.

Summary Male‐biased sex ratios occur in many bird species, particularly in those with small or declining populations, but the causes of these skews and their consequences for local population demography are rarely known. Within‐species variation in sex ratios can help to identify the demographic and behavioural processes associated with such biases.Small populations may be more likely to have skewed sex ratios if sex differences in survival, recruitment or dispersal vary with local abundance. Analyses of species with highly variable local abundances can help to identify these mechanisms and the implications for spatial variation in demography. Many migratory bird species are currently undergoing rapid and severe declines in abundance in parts of their breeding ranges and thus have sufficient spatial variation in abundance to explore the extent of sex ratio biases, their causes and implications.Using national‐scale bird ringing data for one such species (willow warbler, Phylloscopus trochilus), we show that sex ratios vary greatly across Britain and that male‐biased sites are more frequent in areas of low abundance, which are now widespread across much of south and east England. These sex ratio biases are sufficient to impact local productivity, as the relative number of juveniles caught at survey sites declines significantly with increasing sex ratio skew.Sex differences in survival could influence this sex ratio variation, but we find little evidence for sex differences in survival increasing with sex ratio skew. In addition, sex ratios have become male‐biased over the last two decades, but there are no such trends in adult survival rates for males or females. This suggests that lower female recruitment into low abundance sites is contributing to these skews.These findings suggest that male‐biased sex ratios in small and declining populations can arise through local‐scale sex differences in survival and dispersal, with females recruiting disproportionately into larger populations. Given the high level of spatial variation in population declines and abundance of many migratory bird species across Europe at present, male‐biased small populations may be increasingly common. As singing males are the primary records used in surveys of these species, and as unpaired males often sing throughout the breeding season, local sex ratio biases could also be masking the true extent of these population declines.

Male-Biased Capture in Light Traps in Spodoptera exigua (Lepidoptera: Noctuidae): Results from the Studies of Reproductive Activities

Nocturnal insects show phototactic behavior. Interestingly, males are caught by light traps more often than females in many species. In the present study, the male-biased capture in light traps in the field was evaluated to test the hypothesis that male-biased capture was associated with the reproductive activities of Spodoptera exigua. During a 16-wk. period in 2011, a total of 5075 moths caught by light traps in the field revealed a significant male-biased capture. However, the sex ratio of the natural population was not significantly different from 1:1 from July to October 2011. The ratio of moths caught per hour by light traps showed that females were captured in the first half of the night, whereas the peak capture by light traps for males occurred in the second half of the night. Among the captured adults, the percentage of unmated females was significantly lower than that of the mated females and males. Experiments in the laboratory indicated that the time of activities for both sexes was consistent with the active patterns of oviposition and copulation. The percentage of flight-to-light of the unmated females was significantly lower than that of the mated females and males. Thus, the lower capture of the unmated females, which is associated with reduced positive phototactic responses of S. exigua, probably is one of the reasons resulting in the male-biased sex ratio in light traps.