Field estimates of parentage reveal sexually antagonistic selection on body size in a population of Anolis lizards

Sexual dimorphism evolves when selection favors different phenotypic optima between the sexes. Such sexually antagonistic selection creates intralocus sexual conflict when traits are genetically correlated between the sexes and have sex‐specific optima. Brown anoles are highly sexually dimorphic: Males are on average 30% longer than females and 150% heavier in our study population. Viability selection on body size is known to be sexually antagonistic, and directional selection favors large male size whereas stabilizing selection constrains females to remain small. We build on previous studies of viability selection by measuring sexually antagonistic selection using reproductive components of fitness over three generations in a natural population of brown anoles. We estimated the number of offspring produced by an individual that survived to sexual maturity (termed RS^V), a measure of individual fitness that includes aspects of both individual reproductive success and offspring survival. We found directional selection on male body size, consistent with previous studies of viability selection. However, selection on female body size varied among years, and included periods of positive directional selection, quadratic stabilizing selection, and no selection. Selection acts differently in the sexes based on both survival and reproduction and sexual conflict appears to be a persistent force in this species.     

SEASONAL VARIATION IN SEXUAL SELECTION IN THE MOTTLED SCULPIN

Seasonal variation in sexual and natural selection in male mottled sculpins (Cottus bairdi) can be evaluated by calculating selection differentials, which measure the magnitude of phenotypic change resulting from selection, and by calculating indices of the opportunity for selection, which indicate the potential for phenotypic selection in a given interval. Selection differentials are high at the beginning of the breeding season and decline throughout the breeding season. The magnitude and direction of selection differentials depend on when spawning occurs and are independent of the size or age of the females that spawn. Annual selection differentials due to differences in mating success (female choice) are nearly constant between years. Annual selection differentials associated with hatching success are variable. Opportunities for selection (I = fitness variance/[mean fitness]²) show clear seasonal patterns. They are highest at the beginning and at the end of the spawning season. However, this variation is dependent on the mean used to calculate I, and hence variation in I values does not indicate a significant change in the variance of male fitness.     

Life-History Characteristics and Fitness in Descendents of Parthenogenetic and Ex-Ephippio Females of Daphnia Magna

Daphnids of ex-ephippial and parthenogenetic origin differ substantially in life-history. Possible maternally transmitted ex-diapause effects of differing female origin on the fitness of their offspring were studied across multiple clones in Daphnia magna. Ex-ephippial daphnids responded in egg size to different constant food concentrations with the same pattern as parthenogenetic females. Significant differences in egg characteristics between females of different origin were only found for the first clutch produced under high food. The smaller size of eggs in ex-ephippial females, however, did not translate into size differences of first-clutch neonates. A trend for lower mass density per body volume was detected in offspring from ex-ephippial females in comparison with parthenogenetic daphnids. Hence, an ex-diapause effect transmitted by ex-ephippial mothers to their parthenogenetic offspring is likely. However, there was no difference in life-histories and fitness between offspring produced by females of ex-ephippial and parthenogenetic origin at both, high and low food concentrations across multiple clones. Thus, a significant ex-diapause effect on fitness in successive parthenogenetic generations may not be expected in D. magna at the population level.     

SEX-RATIO SELECTION IN A BIVOLTINE THRIPS. I. CONDITIONAL SEX-RATIO MANIPULATION AND FITNESS VARIATION

Females of the bivoltine thrips Elaphrothrips tuberculatus (Hood) (Insecta: Thysanoptera) produce broods of either all males (by viviparity) or all females (by oviparity). Measurements of the sex-allocation ratio, ecological and physiological conditions affecting male and female offspring body size, and correlates of the relative fitnesses of adult males and females in relation to size indicate that female parents tend to be viviparous (produce males) if their offspring will become relatively large adults, and that males gain more in fitness from large size than do females. However, the conditions that link sex allocation with offspring fitness differ between the spring and summer generations. In spring, when breeding is synchronous, 1) oviparous and viviparous females do not differ in body size, 2) females tend to be viviparous where the fungus upon which they feed is relatively dense and where their offspring will become relatively large adults, and 3) fungus density is highly correlated with male and female offspring size. In summer, when breeding is relatively asynchronous, 1) viviparous females are much larger than oviparous females early (but not late) in the season, 2) large viviparous females begin breeding earlier than smaller ones, 3) offspring developing earlier in the season become larger adults, and 4) a higher proportion of females are viviparous earlier than later. Field experiments and field collections show that the covariation among sex allocation, conditions, and fitness is not caused by differential mortality by size or sex. Differences between the spring and summer generations in the cues used by females to adjust offspring sex ratio may be caused by seasonal variation in the factors that affect offspring size. However, in both generations, females tend to produce sons only when their offspring will become relatively large adults, whereas daughters are produced regardless of offspring size. These data suggest that females of E. tuberculatus avoid production of males (the sex with higher variance in expected fitness) when the size of their offspring is relatively uncertain.     

The song of the old mother: Reproductive senescence in female drosophila

Among animals with multiple reproductive episodes, changes in adult condition over time can have profound effects on lifetime reproductive fitness and offspring performance. The changes in condition associated with senescence can be particularly acute for females who support reproductive processes from oogenesis through fertilization. The pomace fly Drosophila melanogaster is a well-established model system for exploring the physiology of reproduction and senescence. In this review, we describe how increasing maternal age in Drosophila affects reproductive fitness and offspring performance as well as the genetic foundation of these effects. Describing the processes underlying female reproductive senescence helps us understand diverse phenomena including population demographics, condition-dependent selection, sexual conflict, and transgenerational effects of maternal condition on offspring fitness. Understanding the genetic basis of reproductive senescence clarifies the nature of life-history trade-offs as well as potential ways to augment and/or limit female fertility in a variety of organisms.     

Rank-Related Fitness Differences and Their Demographic Pathways in Semi-Free-Ranging Rhesus Macaques (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

Researchers have explored the fitness consequences of female dominance hierarchies in many primate populations, with most studies highlighting differences in age of maturation, fertility, and offspring survival. We use resampling techniques and van Tienderen’s (2000) elasticity path analysis to identify rank-related differences in finite rate of increase (λ) and their demographic correlates among segments of a semi-free-ranging rhesus macaque population. Higher-ranking population segments grew at greater rates for some portions of the 40-yr study period. The female members of these segments achieved these lifetime fitness differences through higher fertility and especially higher adult survival rates. This is the first clear evidence that social rank influences female primate adult survival, and is a crucial fitness component for any long-lived, slow-reproducing animal. Traditional methods of comparing lifespans, and other life history variables, among rank categories fail to identify most of the rank-related differences primarily because they require completed life histories that are available only on a small number of the females known in the population.     

Benefits of polyandry: Molecular evidence from field‐caught dung beetles

When females mate with multiple males, they set the stage for postcopulatory sexual selection via sperm competition and/or cryptic female choice. Surprisingly little is known about the rates of multiple mating by females in the wild, despite the importance of this information in understanding the potential for postcopulatory sexual selection to drive the evolution of reproductive behaviour, morphology and physiology. Dung beetles in the genus Onthophagus have become a laboratory model for studying pre‐ and postcopulatory sexual selection, yet we still lack information about the reproductive behaviour of female dung beetles in natural populations. Here, we develop microsatellite markers for Onthophagus taurus and use them to genotype the offspring of wild‐caught females and to estimate natural rates of multiple mating and patterns of sperm utilization. We found that O. taurus females are highly polyandrous: 88% of females produced clutches sired by at least two males, and 5% produced clutches with as many as five sires. Several females (23%) produced clutches with significant paternity skew, indicating the potential for strong postcopulatory sexual selection in natural populations. There were also strong positive correlations between the number of offspring produced and both number of fathers and paternity skew, which suggests that females benefit from mating polyandrously by inciting postcopulatory mechanisms that bias paternity towards males that can sire more viable offspring. This study evaluates the fitness consequences of polyandry for an insect in the wild and provides strong evidence that female dung beetles benefit from multiple mating under natural conditions.     

Female reproductive competition within families in rural Gambia

Many studies show that the extended human family can be helpful in raising offspring, with maternal grandmothers, in particular, improving offspring survival. However, less attention has been given to competition between female kin and co-residents. It has been argued that reproductive conflict between generations explains the evolution of menopause in cooperatively breeding species where females disperse, and that older females are related to the offspring of younger females through their sons, whereas younger, incoming females are unrelated to older females. This means the pattern of help will be asymmetric, so older females lose in reproductive conflict and become 'sterile helpers'. Here, we seek evidence for female reproductive competition using longitudinal demographic data from a rural Gambian population, and examine when women are helping or harming each other's reproductive success. We find that older women benefit and younger women suffer costs of reproductive competition with women in their compound. But the opposite is found for mothers and daughters; if mother and daughter's reproductive spans overlap, the older woman reduces her reproduction if the younger woman (daughter) reproduces, whereas daughters' fertility is unaffected by their mothers' reproduction. Married daughters are not generally co-resident with their mothers, so we find not only competition effects with co-resident females, but also with daughters who have dispersed. Dispersal varies across human societies, but our results suggest reproductive conflict could be influencing reproductive scheduling whatever the dispersal pattern. A cultural norm of late male marriage reduces paternal grandmother/daughter-in-law reproductive overlap almost to zero in this population. We argue that cultural norms surrounding residence and marriage are themselves cultural adaptations to reduce reproductive conflict between generations in human families.     

Giving your daughters the edge: bequeathing reproductive dominance in a primitively social bee

The fitness associated with behavioural strategies is usually estimated in terms of offspring number and size. However, in group-living animals the reproductive value of offspring may also depend on their social rank. We show here that in an allodapine bee Exoneura robusta, dominant mothers can behaviourally influence their daughters'' reproductive rank by controlling insemination of other potential mothers. In E. robusta, group living is near mandatory and reproductive dominance among female nestmates is determined by order of adult emergence. Nests are single, undivided burrows and the dominant female assumes a guarding position closest to the nest entrance. We show that before the egg-laying period, subordinate females who have been absent from the nest are ''screened'' by the reproductive guard upon attempted re-entry. Those who have been in contact with foreign males are less likely to be granted access back into the nest than those who have been in contact with foreign females or with no bees at all. We argue that by controlling insemination patterns of their nestmates, dominant females ensure that their own daughters eclose first and are therefore more likely to assume dominance in the next generation. This presents a situation where dominance is bequeathed to daughters by behavioural means. The ability of mothers to influence social hierarchies in subsequent generations introduces a fitness component additional to the number and size of offspring produced.     

SEQUENTIAL FEMALE ASSESSMENT DRIVES COMPLEX SEXUAL SELECTION ON BOWER SHAPE IN A CICHLID FISH

In many animals, sexual selection on male traits results from female mate choice decisions made during a sequence of courtship behaviors. We use a bower‐building cichlid fish, Nyassachromis cf. microcephalus, to show how applying standard selection analysis to data on sequential female assessment provides new insights into sexual selection by mate choice. We first show that the cumulative selection differentials confirm previous results suggesting female choice favors males holding large volcano‐shaped sand bowers. The sequential assessment analysis reveals these cumulative differentials are the result of selection acting on different bower dimensions during the courtship sequence; females choose to follow males courting from tall bowers, but choose to engage in premating circling with males holding bowers with large diameter platforms. The approach we present extends standard selection analysis by partitioning the variances of increasingly accurate estimates of male reproductive fitness and is applicable to systems in which sequential female assessment drives sexual selection on male traits.     

The effects of sexual selection on life‐history traits: an experimental study on guppies

Sexual selection is often prevented during captive breeding in order to maximize effective population size and retain genetic diversity. However, enforcing monogamy and thereby preventing sexual selection may affect population fitness either negatively by preventing the purging of deleterious mutations or positively by reducing sexual conflicts. To better understand the effect of sexual selection on the fitness of small populations, we compared components of female fitness and the expression of male secondary sexual characters in 19 experimental populations of guppies (Poecilia reticulata) maintained under polygamous or monogamous mating regimes over nine generations. In order to generate treatments that solely differed by their level of sexual selection, the middle‐class neighbourhood breeding design was enforced in the monogamous populations, while in the polygamous populations, all females contributed similarly to the next generation with one male and one female offspring. This experimental design allowed potential sexual conflicts to increase in the polygamous populations because selection could not operate on adult‐female traits. Clutch size and offspring survival showed a weak decline from generation to generation but did not differ among treatments. Offspring size, however, declined across generations, but more in monogamous than polygamous populations. By generation eight, orange‐ and black‐spot areas were larger in males from the polygamous treatment, but these differences were not statistically significant. Overall, these results suggest that neither sexual conflict nor the purging of deleterious mutation had important effects on the fitness of our experimental populations. However, only few generations of enforced monogamy in a benign environment were sufficient to negatively affect offspring size, a trait potentially crucial for survival in the wild. Sexual selection may therefore, under certain circumstances, be beneficial over enforced monogamy during captive breeding.     

The song of the old mother: Reproductive senescence in female drosophila

Among animals with multiple reproductive episodes, changes in adult condition over time can have profound effects on lifetime reproductive fitness and offspring performance. The changes in condition associated with senescence can be particularly acute for females who support reproductive processes from oogenesis through fertilization. The pomace fly Drosophila melanogaster is a well-established model system for exploring the physiology of reproduction and senescence. In this review, we describe how increasing maternal age in Drosophila affects reproductive fitness and offspring performance as well as the genetic foundation of these effects. Describing the processes underlying female reproductive senescence helps us understand diverse phenomena including population demographics, condition-dependent selection, sexual conflict, and transgenerational effects of maternal condition on offspring fitness. Understanding the genetic basis of reproductive senescence clarifies the nature of life-history trade-offs as well as potential ways to augment and/or limit female fertility in a variety of organisms.     

Mate choice for nonadditive genetic benefits and the maintenance of genetic diversity in song sparrows

The lek paradox asserts that strong directional selection via female choice should deplete additive genetic variation in fitness and consequently any benefit to females expressing the preference. Recently, we have provided a novel resolution to the paradox by showing that nonadditive genetic effects such as overdominance can be inherited from parent to offspring, and populations with females that express a mating preference for outbred males maintain higher genetic variation than populations with females that mate randomly. Here, we test our dynamic model using empirical data previously published from a small island population of song sparrows (Melospiza melodia). The model assumes that fitness and male trait expression display overdominance effects. The results demonstrate that female choice for outbred males mediated by directional selection on song repertoire size provides a heritable benefit to offspring through reduced inbreeding depression. Within the population, we estimate the heritability of the inbreeding coefficient to be 0.18 ± 0.08 (SD). Furthermore, we show that mate choice for outbred males increases fitness‐related genetic variation in the population by 12% and thereby reduces inbreeding depression by 1% per generation in typical years and upwards of 15% in severe years. Thus, mate choice may help to stave off population extinction in this and other small populations.     

STABILIZING SELECTION AND VARIANCE IN FIG WASP SEX RATIOS

Theory predicts that the phenotypic variance observed in a trait subject to stabilizing selection should be negatively correlated with the trait's impact on fitness. However, this relationship has rarely been tested directly. The offspring sex ratios produced by pollinating fig wasp foundresses upon entrance to a fruit and oviposition alone (single foundress sex ratios) are subject to stabilizing selection because too many males reduce the total number of dispersing females and too few males will result in unmated females or complete loss of the brood. Furthermore, we argue that the impact on fitness of, and therefore the intensity of stabilizing intensity on, single foundress sex ratios are correlated to how frequently a species produces single foundress broods in nature. Specifically, the intensity of stabilizing selection will be greater in species that encounter single foundress broods more frequently, both because the trait is expressed more often and because fitness shows a greater sensitivity to variation (narrower fitness profile) when that trait is expressed. Across 16 species of Panamanian pollinating fig wasps, the phenotypic variance in single foundress sex ratios was negatively correlated with the frequency with which that species encounters single foundress broods in nature. In addition, a formal comparative analysis based upon a molecular phylogeny of the wasps gave results that were the same as when species were used as independent data points.     

Male house mice evolving with post-copulatory sexual selection sire embryos with increased viability

Although mating is costly, multiple mating by females is a taxonomically widespread phenomenon. Theory has suggested that polyandry may allow females to gain genetic benefits for their offspring, and thus offset the costs associated with this mating strategy. For example, the good sperm hypothesis posits that females benefit from mating multiply when genetically superior males have increased success in sperm competition and produce high quality offspring. We applied the powerful approach of experimental evolution to explore the potential for polyandry to drive evolutionary increases in female fitness in house mice, Mus domesticus. We maintained polygamously mated and monogamously mated selection lines of house mice for 14 generations, before determining whether selection history could account for divergence in embryo viability. We found that males from lineages evolving with post-copulatory sexual selection sire offspring with increased viability, suggesting that polyandry results in the production of higher quality offspring and thus provides long-term fitness benefits to females.     

Positive genetic correlation between female preference and offspring fitness

In many species, females display preferences for extreme male signal traits, but it has not been determined if such preferences evolve as a consequence of females gaining genetic benefits from exercising choice. If females prefer extreme male traits because they indicate male genetic quality that will enhance the fitness of offspring, a genetic correlation will evolve between female preference genes and genes that confer offspring fitness. We show that females of Drosophila serrata prefer extreme male cuticular hydrocarbon (CHC) blends, and that this preference affects offspring fitness. Female preference is positively genetically correlated with offspring fitness, indicating that females have gained genetic benefits from their choice of males. Despite male CHCs experiencing strong sexual selection, the genes underlying attractive CHCs also conferred lower offspring fitness, suggesting a balance between sexual selection and natural selection may have been reached in this population.     

FEMALES ARE CHOOSIER IN THE DARK: ENVIRONMENT‐DEPENDENT RELIANCE ON COURTSHIP COMPONENTS AND ITS IMPACT ON FITNESS

A broad understanding of multimodal courtship function necessitates knowledge of the potential information content of signal components, the efficacy of signal components in eliciting the appropriate receiver response, and the fitness consequences of mating decisions based upon various signal components. We present data addressing each of these requirements for the multimodal‐signaling wolf spider, Schizocosa floridana Bryant. Using diet manipulations, we first demonstrate that both visual and seismic courtship signals are condition‐dependent. Next, using high‐ and low‐quantity diet individuals in mate choice trials across manipulated signaling environments, we demonstrate that the seismic signal is crucial for mating success and further show that female choosiness is environment‐dependent. Females mated more with high diet males only in the absence of visual signals, showing no discrimination in the presence of visual signals. Finally, by quantifying the number of offspring produced by our mated females, we reveal that a female's mating environment, in conjunction with her potential resource availability, influences her fitness—in environments in which females exerted choice, heavier females produced more offspring. Together, this comprehensive set of experiments demonstrates that female choosiness varies across environments, leading to direct fitness consequences.     

Males' evolutionary responses to experimental removal of sexual selection

We evaluated the influence of pre- and post-copulatory sexual selection upon male reproductive traits in a naturally promiscuous species, Drosophila melanogaster. Sexual selection was removed in two replicate populations through enforced monogamous mating with random mate assignment or retained in polyandrous controls. Monogamous mating eliminates all opportunities for mate competition, mate discrimination, sperm competition, cryptic female choice and, hence, sexual conflict. Levels of divergence between lines in sperm production and male fitness traits were quantified after 38-81 generations of selection. Three a priori predictions were tested: (i) male investment in spermatogenesis will be lower in monogamy-line males due to the absence of sperm competition selection, (ii) due to the evolution of increased male benevolence, the fitness of females paired with monogamy-line males will be higher than that of females paired with control-line males, and (iii) monogamy-line males will exhibit decreased competitive reproductive success relative to control-line males. The first two predictions were supported, whereas the third prediction was not. Monogamy males evolved a smaller body size and the size of their testes and the number of sperm within the testes were disproportionately further reduced. In contrast, the fitness of monogamous males (and their mates) was greater when reproducing in a non-competitive context: females mated once with monogamous males produced offspring at a faster rate and produced a greater total number of surviving progeny than did females mated to control males. The results indicate that sexual selection favours the production of increased numbers of sperm in D. melanogaster and that sexual selection favours some male traits conferring a direct cost to the fecundity of females.     

Social and genetic benefits of parental investment suggest sex differences in selection pressures

Provisioning behavior in altricial birds is often used to measure parental investment and is assumed to have fitness consequences to the parents providing it, with the benefits outweighing the costs. Here we investigate the fitness costs and benefits (parent survival and offspring recruitment) of provisioning behavior in wild house sparrows Passer domesticus, using long‐term data from a pedigreed isolated population. We disentangled the long‐term fitness consequences in terms of number of recruits, of provisioning behavior from those of other parental investments and individual quality through a cross‐foster design. We accounted for extra‐pair offspring in all analyses. Provisioning behavior confers social fitness benefits in terms of the number of recruits to both parents. Only in females we detected an influence individual quality: female sparrows with high provisioning frequencies were associated with more genetic recruits than those who provided food less frequently to their young, even though foster parents reared the offspring. We detected a relationship between annual survival probability and provisioning behavior only in males, but not in females. This finding, together with indirect benefits differing by sex, indicates that different selection pressures are acting on the sexes. Our study can show that it is justified to use provisioning behavior as a form of parental investment sensu Trivers, since we show that this behavior is costly to parents and that the genetic fitness benefits exceed the costs.     

Effects of exposure to short-term heat stress on fitness components in Drosophila melanogaster

Effects of thermal stress on survival and reproductive success in ten recently collected isofemale lines of Drosophila melanogaster were compared for flies treated as follows: always held at 25° C, placed in an incubator set at 37° C for 120 min, or exposed to 40° C in an incubator for 90 min, with or without previous exposure to 37° C. Short-term exposure to the higher temperature greatly reduced adult survival, the mating frequency of males and females, and female fecundity, which was measured as offspring produced over ten days. Male fertility, measured as the progeny produced by a female mated once, differed little among treatments. Previous exposure to a high, but non-lethal, temperature before exposure to the higher one, improved survival of males and females, and improved offspring production of females. Genetic variation was present among lines for offspring production, but genetic variation for survival was not significant, and genotype by environment interactions for fitness components of females were small. These results indicated low genetic variation in thermal resistance in the studied population, such that a threshold for temperature stress probably exists, above which local extinction is more likely than the evolution of resistance.