Clonal reproduction by males and females in the little fire ant

Sexual reproduction can lead to major conflicts between sexes and within genomes. Here we report an extreme case of such conflicts in the little fire ant Wasmannia auropunctata. We found that sterile workers are produced by normal sexual reproduction, whereas daughter queens are invariably clonally produced. Because males usually develop from unfertilized maternal eggs in ants and other haplodiploid species, they normally achieve direct fitness only through diploid female offspring. Hence, although the clonal production of queens increases the queen's relatedness to reproductive daughters, it potentially reduces male reproductive success to zero. In an apparent response to this conflict between sexes, genetic analyses reveal that males reproduce clonally, most likely by eliminating the maternal half of the genome in diploid eggs. As a result, all sons have nuclear genomes identical to those of their father. The obligate clonal production of males and queens from individuals of the same sex effectively results in a complete separation of the male and female gene pools. These findings show that the haplodiploid sex-determination system provides grounds for the evolution of extraordinary genetic systems and new types of sexual conflict.     

Negative genetic correlation between male sexual attractiveness and survival

Indirect selection of female mating preferences may result from a genetic association between male attractiveness and offspring fitness. The offspring of attractive males may have enhanced growth, fecundity, viability or attractiveness. However, the extent to which attractive males bear genes that reduce other fitness components has remained unexplored. Here I show that sexual attractiveness in male guppies (Poecilia reticulata) is heritable and genetically correlated with ornamentation. Like ornamentation, attractiveness may be substantially Y-linked. The benefit of mating with attractive males, and thus having attractive sons, is opposed by strong negative genetic correlation between attractiveness and both offspring survival and the number of sons maturing. Such correlations suggest either antagonistic pleiotropy between attractiveness and survival or linkage disequilibrium between attractive and deleterious alleles. The presence of many colour pattern genes on or near the non-recombining section of the Y chromosome may facilitate the accumulation of deleterious mutations by genetic hitchhiking. These findings show that genes enhancing sexual attractiveness may be associated with pleiotropic costs or heavy mutational loads.     

The adaptive significance of temperature-dependent sex determination in a reptile

Understanding the mechanisms that determine an individual's sex remains a primary challenge for evolutionary biology. Chromosome-based systems (genotypic sex determination) that generate roughly equal numbers of sons and daughters accord with theory, but the adaptive significance of environmental sex determination (that is, when embryonic environmental conditions determine offspring sex, ESD) is a major unsolved problem. Theoretical models predict that selection should favour ESD over genotypic sex determination when the developmental environment differentially influences male versus female fitness (that is, the Charnov-Bull model), but empirical evidence for this hypothesis remains elusive in amniote vertebrates--the clade in which ESD is most prevalent. Here we provide the first substantial empirical support for this model by showing that incubation temperatures influence reproductive success of males differently than that of females in a short-lived lizard (Amphibolurus muricatus, Agamidae) with temperature-dependent sex determination. We incubated eggs at a variety of temperatures, and de-confounded sex and incubation temperature by using hormonal manipulations to embryos. We then raised lizards in field enclosures and quantified their lifetime reproductive success. Incubation temperature affected reproductive success differently in males versus females in exactly the way predicted by theory: the fitness of each sex was maximized by the incubation temperature that produces that sex. Our results provide unequivocal empirical support for the Charnov-Bull model for the adaptive significance of temperature-dependent sex determination in amniote vertebrates.     

Population density affects sex ratio variation in red deer.

Many mammal populations show significant deviations from an equal sex ratio at birth, but these effects are notoriously inconsistent. This may be because more than one mechanism affects the sex ratio and the action of these mechanisms depends on environmental conditions. Here we show that the adaptive relationship between maternal dominance and offspring sex ratio previously demonstrated in red deer (Cervus elaphus), where dominant females produced more males, disappeared at high population density. The proportion of males born each year declined with increasing population density and with winter rainfall, both of which are environmental variables associated with nutritional stress during pregnancy. These changes in the sex ratio corresponded to reductions in fecundity, suggesting that they were caused by differential fetal loss. In contrast, the earlier association with maternal dominance is presumed to have been generated pre-implantation. The effects of one source of variation superseded the other within about two generations. Comparison with other ungulate studies indicates that positive associations between maternal quality and the proportion of male offspring born have only been documented in populations below carrying capacity.     

Sexually antagonistic genetic variation for fitness in red deer

Evolutionary theory predicts the depletion of genetic variation in natural populations as a result of the effects of selection, but genetic variation is nevertheless abundant for many traits that are under directional or stabilizing selection. Evolutionary geneticists commonly try to explain this paradox with mechanisms that lead to a balance between mutation and selection. However, theoretical predictions of equilibrium genetic variance under mutation-selection balance are usually lower than the observed values, and the reason for this is unknown. The potential role of sexually antagonistic selection in maintaining genetic variation has received little attention in this debate, surprisingly given its potential ubiquity in dioecious organisms. At fitness-related loci, a given genotype may be selected in opposite directions in the two sexes. Such sexually antagonistic selection will reduce the otherwise-expected positive genetic correlation between male and female fitness. Both theory and experimental data suggest that males and females of the same species may have divergent genetic optima, but supporting data from wild populations are still scarce. Here we present evidence for sexually antagonistic fitness variation in a natural population, using data from a long-term study of red deer (Cervus elaphus). We show that male red deer with relatively high fitness fathered, on average, daughters with relatively low fitness. This was due to a negative genetic correlation between estimates of fitness in males and females. In particular, we show that selection favours males that carry low breeding values for female fitness. Our results demonstrate that sexually antagonistic selection can lead to a trade-off between the optimal genotypes for males and females; this mechanism will have profound effects on the operation of selection and the maintenance of genetic variation in natural populations.     

Fitness costs of gestation and lactation in wild mammals

Like a number of plants, some mammals commonly produce more progeny than they can afford to rear, terminating investment in some or even all of their offspring once the resources available for breeding are known. Adaptive interpretations of juvenile wastage rely on the argument that the costs of gestation are small compared to those of feeding offspring. Though energetic evidence supports this conclusion, it is unsafe to assume that the relative costs of gestation and lactation to the mother's survival and future reproductive success follow the same pattern because lactation commonly coincides with the period of maximum food availability. Controlling for individual variation, we show that in wild red deer (Cervus elaphus L) any costs of gestation to the mother's subsequent survival and reproductive success are slight compared to those of lactation.     

Sex differences in emigration and mortality affect optimal management of deer populations

Populations of red deer that are limited by food, like those of many other ungulates, commonly include more females than males. We assessed the contribution of variation in sex- and age-specific rates of mortality and emigration to density-dependent changes in the adult sex ratio, using long-term observations and demographic experiments involving the red deer population on Rum, Scotland. We incorporated these effects in a stochastic model of local populations under different management regimes to show here that, when female numbers are allowed to increase to more than 60% of the ecological carrying capacity, the sustainable annual harvest of males from local deer populations will fall. Because males are typically culled by fee-paying hunters and generate more income than females, income will decrease as the male harvest falls. Because numbers of female deer throughout much of the Highlands probably exceed the threshold at which male density starts to be affected, many managers might be able to raise income from local deer populations by reducing female numbers, with potential benefits to the vegetation of Scottish Highland environments.     

非损伤技术研究天山马鹿性别比与冬季家域

 利用5对微卫星对331 堆粪便进行个体识别,发现喀拉乌成山小渠子地区73 堆粪便来自32 头马鹿,白杨沟64 堆粪便来自28 头马鹿,哈密东天山国家森林公园的白石头地区87 堆粪便来自66 头马鹿,松树塘地区41 堆粪便来自32 头马鹿,寒气沟地区66 堆粪便来自42 头马鹿。利用SRY基因鉴定的天山马鹿雌雄比是1.78:1~2.89:1,粪便形态鉴定的结果是1.62:1~2.40:1,两种方法获得的结果无统计学差异。冬季雌雄马鹿家域范围分别是1.24~1.71 km²和1.02~1.57 km²,雌雄性家域范围差异不大,这是由天山马鹿自身习性决定的。     

The optimal sex ratio in cooperatively breeding populations

In some cooperatively breeding species, elder siblings remain within their home ranges to assist their parents in raising their younger siblings by providing protection or food. Previous attempts to model the popu- lation sex ratios of such species have assumed that helping is cost-free and only involves benefits to parents, resulting in offspring sex ratio biases towards the helping sex. However, parents may also incur costs in maintaining helpers. We incorporated costs and benefits of helping to predict the sex ratio of offspring in a theoretical model. Our model showed that the evolutionary stable strategy （ESS） sex ratio strongly depends on three parameters： （1） the average number of helpers per female; （2） the benefits that a helper brings to its mother＇s offspring production （MOP）; and （3） the cost-benefit ratio of helping. When one sex of elder siblings provides help, the ESS sex ratio is biased towards the helping sex if MOP costs are less than the benefits, （i.e., the cost-benefit ratio 〈1）. However, the ESS sex ratio becomes biased towards the non-helping sex if MOP costs exceed the benefits （i.e., the cost-benefit ratio 〉1）. Additionally, our model shows that the ESS sex ratio becomes biased in favor of the more helpful sex when both male and female elder siblings provide help. These results explain why sex ratios may become biased towards the helping sex, as well as the non-helping sex, in some species.     

Hybridization and adaptive mate choice in flycatchers

Hybridization in natural populations is strongly selected against when hybrid offspring have reduced fitness. Here we show that, paradoxically, pairing with another species may offer the best fitness return for an individual, despite reduced fitness of hybrid offspring. Two mechanisms reduce the costs to female collared flycatchers of pairing with male pied flycatchers. A large proportion of young are sired by conspecific male collared flycatchers through extra-pair copulations, and there is a bias in favour of male offspring (which, unlike females, are fertile) within hybrid pairs. In combination with temporal variation in breeding success, these cost-reducing mechanisms yield quantitative predictions about when female collared flycatchers should accept a male pied flycatcher as a mate; empirical data agree with these predictions. Apparent hybridization may thus represent adaptive mate choice under some circumstances.     

Rank-related maternal effects of androgens on behaviour in wild spotted hyaenas

Within any hierarchical society, an individual's social rank can have profound effects on its health and reproductive success, and rank-related variation in these traits is often mediated by variation in endocrine function. Maternal effects mediated by prenatal hormone exposure are potentially important for non-genetic inheritance of phenotypic traits related to social rank, and thus for shaping individual variation in behaviour and social structure. Here we show that androgen concentrations in wild female spotted hyaenas (Crocuta crocuta) are higher during late gestation in dominant females than in subordinate females. Furthermore, both male and female cubs born to mothers with high concentrations of androgens in late pregnancy exhibit higher rates of aggression and mounting behaviour than cubs born to mothers with lower androgen concentrations. Both behaviours are strongly affected in other mammals by organizational effects of androgens, and both have important effects on fitness in hyaenas. Therefore, our results suggest that rank-related maternal effects of prenatal androgen exposure can adaptively influence offspring phenotype in mammals, as has previously been shown to occur in birds. They also suggest an organizational mechanism for the development of female dominance and aggressiveness in spotted hyaenas, traits that may offset the costs of extreme virilization.     

Paternal inheritance of a female moth's mating preference

Females of the arctiid moth Utetheisa ornatrix mate preferentially with larger males, receiving both direct phenotypic and indirect genetic benefits. Here we demonstrate that the female's mating preference is inherited through the father rather than the mother, indicating that the preference gene or genes lie mostly or exclusively on the Z sex chromosome, which is strictly paternally inherited by daughters. Furthermore, we show that the preferred male trait and the female preference for that trait are correlated, as females with larger fathers have a stronger preference for larger males. These findings are predicted by the protected invasion theory, which asserts that male homogametic sex chromosome systems (ZZ/ZW) found in lepidopterans and birds promote the evolution of exaggerated male traits through sexual selection. Specifically, the theory predicts that, because female preference alleles arising on the Z chromosome are transmitted to all sons that have the father's attractive trait rather than to only a fraction of the sons, such alleles will experience stronger positive selection and be less vulnerable to chance loss than would autosomal alleles.     

杨万里的儿女姻亲考述

杨万里生四子五女，一子早夭。本文对其儿女生平及其姻亲状况一一作出考述，以就正于通识大家。     

Adaptive variation in environmental and genetic sex determination in a fish

Two general mechanisms of sex determination have been identified among gonochoristic vertebrates: environmental sex determination where offspring become male or female in response to an environmental factor(s) during development (for example, some fishes and reptiles); and genetic sex determination where sex is determined by genotype at conception (as in birds and mammals). How do these sex-determining systems evolve? Direct evidence is virtually non-existent because the sex-determining systems of most species appear to have little genetic variation. Here we provide the first evidence of adaptive variation in environmental and genetic sex determination within a species. We show that in a fish with temperature-dependent sex determination, populations at different latitudes compensate for differences in thermal environment and seasonality by adjusting the response of sex ratio to temperature, and by altering the level of environmental as opposed to genetic control. The adjustments observed are precisely those predicted by adaptive sex ratio theory.     

A socially enforced signal of quality in a paper wasp

Organisms use signals of quality to communicate information about aspects of their relative phenotypic and genetic constitution. Badges of status are a subset of signals of quality that reveal information about an individual's size and dominance. In general, signals of quality require high and differential costs to remain honest (that is, prevent low-quality cheaters from exploiting any fitness benefits associated with communicating high quality). The theoretically required costs for badges of status remain controversial because the development (or 'production') of such signals often seems to be relatively cost-free. One important hypothesis is that such signals impose social (or 'maintenance') costs incurred through repeated agonistic interactions with other individuals. However, convincing empirical evidence for social costs remains elusive. Here we report social costs in a previously undescribed badge of status: the highly variable black facial patterns of female paper wasps, Polistes dominulus. Facial patterns strongly predict body size and social dominance. Moreover, in staged contests between pairs of unfamiliar wasps, subordinate wasps with experimentally altered facial features ('cheaters') received considerably more aggression from the dominant than did sham controls, indicating that facial patterns are signals and that dishonest signalling imposes social costs.     

Female transference and mate choice among Tana River red colobus.

Red colobus are one of a small number of primate species in which females have been reported to transfer between breeding groups more commonly than males. Several authors have hypothesised that in such species transference may serve to reduce the risk to females of producing offspring of lower fitness through inbreeding. The hypothesis offers no explanation of why females rather than males are responsible for outbreeding in these species, but remains plausible so long as male membership of breeding groups is relatively stable; for once members of one sex have evolved dispersal mechanisms reducing the risk of inbreeding, pressures on the other sex to do likewise will be lowered. Hence, if both sexes commonly migrate, the hypothesis is weakened. I describe here the membership dynamics of a group of Tana River red colobus, Colobus badius rufomitratus, which provide the first evidence of high rates of membership turnover by both sexes in primates, and speculate that the function of female transference in this case may be related to mate choice and the avoidance of infanticide.     

Sexual conflict reduces offspring fitness in zebra finches

Parental care is often costly; hence, in sexually reproducing species where both male and female parents rear their offspring (biparental care), sexual conflict over parental investment can arise. Such conflict occurs because each care-giver would benefit from withholding parental investment for use with another partner, leading to a reduction in the amount of care given by one parent at the expense of the other. Here we report experiments to explore the prediction from theory that parents rearing offspring alone may provide greater parental investment than when rearing offspring together with a partner. We found that when the number of offspring per parent, and hence the potential workload, were kept constant, offspring received a greater per capita parental investment from single females than from both parents working together, and that males reared by single mothers were more sexually attractive as adults than their biparentally reared siblings. This difference between single- and two-parent families is due to a reduction in care provided by females when they care together with a male, rather than laziness by males or differences in the begging behaviour of chicks, supporting the claim that sexual conflict in biparental care can reduce the quality of offspring raised.     

Embryonic temperature determines adult sexuality in a reptile

Gonadal differentiation in amniote vertebrates is controlled by one of two mechanisms: genotypic sex determination (GSD) or environmental sex determination (ESD). After differentiation the fetal gonad produces sex steroid hormones which govern the development of other components of sexuality. Thus, the primary sex determiner is thought to operate solely as a trigger that initiates a cascade of events culminating in adult sex differences. In the leopard gecko (Eublepharis macularius), gonadal and morphological sex is determined by incubation temperature, with relatively 'hot' temperatures (32 degrees C) resulting in mostly male offspring and relatively 'cold' temperatures (26 degrees C) resulting in only female offspring. We report here that the reproductive behaviour and endocrine physiology of an adult is influenced by the temperature experienced as an embryo. Also, the perception of a female to courtship by a male is influenced by incubation temperature. These data indicate that incubation temperature, the primary determiner of sex in this species, has differential effects on adult sexuality.     

Female mating bias results in conflicting sex-specific offspring fitness

Indirect-benefit models of sexual selection assert that females gain heritable offspring advantages through a mating bias for males of superior genetic quality. This has generally been tested by associating a simple morphological quality indicator (for example, bird tail length) with offspring viability. However, selection acts simultaneously on many characters, limiting the ability to detect significant associations, especially if the simple indicator is weakly correlated to male fitness. Furthermore, recent conceptual developments suggest that the benefits gained from such mating biases may be sex-specific because of sexually antagonistic genes that differentially influence male and female reproductive ability. A more suitable test of the indirect-benefit model would examine associations between an aggregate quality indicator (such as male mating success) and gender-specific adult fitness components, under the expectation that these components may trade off. Here, we show that a father's mating success in the cricket, Allonemobius socius, is positively genetically correlated with his son's mating success but negatively with his daughter's reproductive success. This provides empirical evidence that a female mating bias can result in sexually antagonistic offspring fitness.     

Intrasexual competition and sexual selection in cooperative mammals

In most animals, the sex that invests least in its offspring competes more intensely for access to the opposite sex and shows greater development of secondary sexual characters than the sex that invests most. However, in some mammals where females are the primary care-givers, females compete more frequently or intensely with each other than males. A possible explanation is that, in these species, the resources necessary for successful female reproduction are heavily concentrated and intrasexual competition for breeding opportunities is more intense among females than among males. Intrasexual competition between females is likely to be particularly intense in cooperative breeders where a single female monopolizes reproduction in each group. Here, we use data from a twelve-year study of wild meerkats (Suricata suricatta), where females show high levels of reproductive skew, to show that females gain greater benefits from acquiring dominant status than males and traits that increase competitive ability exert a stronger influence on their breeding success. Females that acquire dominant status also develop a suite of morphological, physiological and behavioural characteristics that help them to control other group members. Our results show that sex differences in parental investment are not the only mechanism capable of generating sex differences in reproductive competition and emphasize the extent to which competition for breeding opportunities between females can affect the evolution of sex differences and the operation of sexual selection.