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Density-related changes in sexual selection in red deer.

In sexually dimorphic mammals, high population density is commonly associated with increased mortality of males relative to females and with female-biased adult sex ratios. This paper investigates the consequences of these changes on the distribution of male breeding success, the intensity of competition for females and the opportunity for sexual selection. After the red deer (Cervus elaphus L.) population of the North Block of Rum (Inner Hebrides) was released from culling, female numbers rose and male numbers declined, leading to an adult sex ratio of around one male to two females. This change was the result of increased mortality of males relative to females during the first two years of life; of increased emigration rates by young males; and of reduced immigration by males from outside the study area. The increasing bias in the adult sex ratio affected the timing of breeding as well as the distribution of mating success in males. As the adult sex ratio became increasingly biased towards females, the degree of skew in mating success (calculated across all harem-holders) increased, but mature males defended harems for shorter periods and a higher proportion of males held harems. In addition, a higher proportion of calves were fathered by immigrant males and the proportion fathered by males born in the study area declined. These results support the contention that, where high population density is associated with a female-biased adult sex ratio, competition for mates is likely to decline.

Animals↗

Orientation of the genetic variance-covariance matrix and the fitness surface for multiple male sexually selected traits.

Stabilizing selection has been predicted to change genetic variances and covariances so that the orientation of the genetic variance-covariance matrix (G) becomes aligned with the orientation of the fitness surface, but it is less clear how directional selection may change G. Here we develop statistical approaches to the comparison of G with vectors of linear and nonlinear selection. We apply these approaches to a set of male sexually selected cuticular hydrocarbons (CHCs) of Drosophila serrata. Even though male CHCs displayed substantial additive genetic variance, more than 99% of the genetic variance was orientated 74.9 degrees away from the vector of linear sexual selection, suggesting that open-ended female preferences may greatly reduce genetic variation in male display traits. Although the orientation of G and the fitness surface were found to differ significantly, the similarity present in eigenstructure was a consequence of traits under weak linear selection and strong nonlinear (convex) selection. Associating the eigenstructure of G with vectors of linear and nonlinear selection may provide a way of determining what long-term changes in G may be generated by the processes of natural and sexual selection.

Animals↗

Direct and indirect sexual selection and quantitative genetics of male traits in guppies (Poecilia reticulata).

The ornamentation and displays on which sexual attractiveness and thus mating success are based may be complex and comprise several traits. Predicting the outcome of sexual selection on such complex phenotypes requires an understanding of both the direct operation of selection on each trait and the indirect consequences of selection operating directly on genetically correlated traits. Here we report the results of a quantitative genetic analysis of the ornamentation, sexual attractiveness, and mating success of male guppies (Poecilia reticulata). We analyze male ornamentation both from the point of view of single ornamental traits (e.g., the area of each color) and of composite measures of the way the entire pattern is likely to be perceived by females (e.g., the mean and contrast in chroma). We demonstrate that there is substantial additive genetic variation in almost all measures of male ornamentation and that much of this variation may be Y linked. Attractiveness and mating success are positively correlated at the phenotypic and genetic level. Orange area and chroma, the area of a male's tail, and the color contrast of his pattern overall are positively correlated with attractiveness and/or mating success at the phenotypic and genetic levels. Using attractiveness and mating success as measures of fitness, we estimate gradients of linear directional sexual selection operating on each male trait and use equations of multivariate evolutionary change to predict the response of male ornamentation to this sexual selection. From these analyses, we predict that indirect selection may have important effects on the evolution of male guppy color patterns.

Animals↗

Nest building is a sexually selected behaviour in the barn swallow.

Females may use male nest building to assess male parental quality, and nest size would then be a sexually selected trait. In the barn swallow, Hirundo rustica, females select their partner by his tail length, a character believed to signal good genes. Both sexes participate in nest building, although male participation is negatively related to his attractiveness as reflected by tail length. We tested the hypothesis that nest building is a sexually selected trait: females paired with males of high parental quality (as shown by the male during nest building) may obtain a mate providing large amounts of parental investment, while, as has been shown previously, females mated to attractive (long-tailed) males will acquire mates with good genetic quality. Therefore, since nest building in barn swallows occurs after mating, we predicted a postmating sexual selection process by which the female invests differentially in reproduction depending on the male's nest-building effort (reflecting his willingness to invest in reproduction). The volume of material in a nest was related to the male's contribution to nest building and, in agreement with our hypothesis, in a multiple regression analysis, male tail length and nest material volume were negatively related to laying date and positively to female investment in reproduction (total number of eggs laid during the breeding season). Moreover, females paired with long-tailed males (which contribute very little to nest building), but using the same amount of nest material as females paired with short-tailed males, reduced the thickness of the nest and hence increased its capacity. Therefore, in the barn swallow two different traits appear to be sexually selected: tail length of males owing to the good genes process and nest-building ability owing to the good parent process. (c) 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

How do natural and sexual selection contribute to sympatric speciation?

I use explicit genetic models to investigate the importance of natural and sexual selection during sympatric speciation and to sort out how genetic architecture influences these processes. Assortative mating alone can lead to speciation, but rare phenotypes' disadvantage in finding mates and intermediate phenotypes' advantage due to stabilizing selection strongly impede speciation. Any increase in the number of loci also decreases the likelihood of speciation. Sympatric speciation is then harder to achieve than previously demonstrated by many theoretical studies which assume no mating disadvantage for rare phenotypes and consider a small number of loci. However, when a high level of assortative mating evolves, sexual selection might allow populations to split into dimorphic distributions with peaks corresponding to nearly extreme phenotypes. Competition then works against speciation by favouring intermediate phenotypes and preventing further divergence. The interplay between natural and sexual selection during speciation is then more complex than previously explained.

Animals↗

Sexual selection by male choice in monogamous and polygynous human populations.

The theoretical possibility of coevolution of a viability-reducing female physical trait and a male mating preference for that trait by Fisherian sexual selection in monogamous and polygynous populations is demonstrated using two-locus haploid models. It is assumed that there is dichotomous variation in male resources, resource-rich males have a wider choice among females than resource-poor males, and a female has greater reproductive success when mated with a resource-rich male than a resource-poor one. Under these assumptions, we find that sexual selection operates effectively when female reproductive success is strongly dependent on male resource, the proportion of females that mate with resource-rich males is neither small nor large, the degree of polygyny is low, and resources are inherited from father to son. We suggest that some human female physical traits may have evolved by sexual selection through male choice. The evolution of skin color by sexual selection is discussed as an example.

Chromosome Mapping↗

Adult fitness consequences of sexual selection in Drosophila melanogaster.

Few experiments have demonstrated a genetic correlation between the process of sexual selection and fitness benefits in offspring, either through female choice or male competition. Those that have looked at the relationship between female choice and offspring fitness have focused on juvenile fitness components, rather than fitness at later stages in the life cycle. In addition, many of these studies have not controlled for possible maternal effects. To test for a relationship between sexual selection and adult fitness, we carried out an artificial selection experiment in the fruit fly, Drosophila melanogaster. We created two treatments that varied in the level of opportunity for sexual selection. Increased opportunity for female choice and male competition was genetically correlated with an increase in adult survivorship, as well as an increase in male and female body size. Contrary to previous, single-generation studies, we did not find an increase in larval competitive ability. This study demonstrates that mate choice and/or male-male competition are correlated with an increase in at least one adult fitness component of offspring.

Aging↗

The impact of sexual selection on Corynosoma magdaleni (Acanthocephala) infrapopulations in Saimaa ringed seals (Phoca hispida saimensis).

In free-living animals sexual selection is a central force shaping the spatial distribution of individuals in a population as well as sexual size dimorphism. We studied the influence of sexual selection on spatial distribution, female-to-male body size ratio, and female mating success of acanthocephalans in a natural host population of Saimaa ringed seal (Phoca hispida saimensis) harbouring a single intestinal helminth species, Corynosoma magdaleni. The acanthocephalans were always found along the full length of the small intestine; however, the site selection varied among the individual seals according to the age of the infection. The distribution of male acanthocephalans was not random with respect to females, with larger males tending to aggregate around non-mated females. A higher proportion of C. magdaleni females had copulated in seals with relatively more male worms. Male-male competition for access to females can be intense in C. magdaleni infrapopulation and may select for large males. We found that the larger the infrapopulation size, the smaller the males compared to females. In addition, the greater the female bias in the infrapopulation, the smaller the testes of males. Our study shows that sexual selection may be an important determinant of spatial distribution, male body size and female mating success of C. magdaleni in Saimaa ringed seal.

Acanthocephala↗

Sexual selection and speciation in hawaiian Drosophila.

Many ideas about sexual selection and speciation have been stimulated by considering Hawaiian Drosophila. In turn, the ideas and models have stimulated research with the flies, particularly members of the planitibia group. This paper describes high points of some of the models and reviews data that are relevant to understanding sexual selection and speciation in this species group.

Animals↗

Elevated testosterone reduces choosiness in female dark-eyed juncos (Junco hyemalis): evidence for a hormonal constraint on sexual selection?

Because testosterone (T) often mediates the expression of attractive displays and ornaments, in the absence of constraints sexual selection should lead to an evolutionary increase in male T levels. One candidate constraint would be a genetic correlation between the sexes that leads to a correlated response in females. If increased T in females were to have deleterious effects on mate choice, the effect of sexual selection on male T would be weakened. Using female dark-eyed juncos (Junco hyemalis), we tested whether experimentally enhancing female T would lead to a decrease in discrimination between two classes of males, one treated with T (T-males) and one control (C-males). The two female treatments (T-implanted and C-females) spent equal amounts of time with both classes of males, but T-treated females failed to show a preference for either male treatment, whereas C-females showed a significant preference, albeit in an unexpected direction (for C-males). T-females were less discriminating than C-females, irrespective of the direction of their preference. To our knowledge, this is the first study to show that circulating hormones can alter female choosiness without reducing sexual motivation. Our results suggest that hormonal correlations between the sexes have the potential to constrain sexual selection on males.

Analysis of Variance↗

Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae).

Both ornaments and weapons of sexual selection frequently exhibit prolific interspecific diversity of form. Yet, most studies of this diversity have focused on ornaments involved with female mate choice, rather than on the weapons of male competition. With few exceptions, the mechanisms of divergence in weapon morphology remain largely unexplored. Here, we characterize the evolutionary radiation of one type of weapon: beetle horns. We use partial sequences from four nuclear and three mitochondrial genes to develop a phylogenetic hypothesis for a worldwide sample of 48 species from the dung beetle genus Onthophagus (Coleoptera: Scarabaeidae). We then use these data to test for multiple evolutionary origins of horns and to characterize the evolutionary radiation of horns. Although our limited sampling of one of the world's most species-rich genera almost certainly underestimates the number of evolutionary events, our phylogeny reveals prolific evolutionary lability of these exaggerated sexually selected weapons (more than 25 separate gains and losses of five different horn types). We discuss these results in the context of the natural history of these beetles and explore ways that sexual selection and ecology may have interacted to generate this extraordinary diversity of weapon morphology.

Adaptation, Biological↗

Estimating the strength of sexual selection from Y-chromosome and mitochondrial DNA diversity.

We show that a sex difference in the opportunity for selection results in sex differences in the strength of random genetic drift and thus creates different patterns of genetic diversity for maternally and paternally inherited haploid genes. We derive the effective population size Ne for a male-limited or female-limited haploid gene in terms of I, the "opportunity for selection" or the variance in relative fitness. Because the variance in relative fitness of males can be an order of magnitude larger than that of females, the Ne is much smaller for males than it is for females. We derive both nonequilibrium and equilibrium expressions for F(ST) in terms of I and show how the portion of I owing to sexual selection, Imates, that is, the variation among males in mate numbers, is a simple function of the F's for cytoplasmic (female inherited) and Y-linked (male inherited) genes. Because multiple, transgenerational data are lacking to apply the nonequilibrium expression, we apply only the equilibrium model to published data on Y chromosome and mitochondrial sequence divergence in Homo sapiens to quantify the opportunity for sexual selection. The estimate suggests that sexual selection in humans represents a minimum of 54.8% of total selection, supporting Darwin's proposal that sexual selection has played a significant role in human evolution and the recent proposal regarding a shift from polygamy to monogamy in humans.

Chromosomes, Human, Y↗

Domesticated birds as a model for the genetics of speciation by sexual selection.

In theory, even populations occupying identical environments can diverge in sexually selected traits, as a consequence of different mutational input. I evaluate the potential of this process by comparing the genetics of breeds of domesticated birds to what is known about the genetics of differences among species. Within domesticated species there is a strong correlation of time since domestication with the number of breeds. Descendants of the rock dove, Columba livia (the oldest domesticate) show differences in courtship, vocalizations, body shape, feather ornaments (crests and tails) and colors and color patterns. When nine other domesticated species are included there is a striking hierarchy, with more recent domesticates having a nested subset of these traits: the youngest domesticated species have breeds distinguished only by color. This suggests that selection of new, visible, mutations is driving the process of breed diversification, with mutations that appeal to the breeder happening the most frequently in color. In crosses among related species, color, feather ornaments and many vocalizations and displays show both intermediate dominance and pure dominance. Although the number of loci affecting each of these traits is typically unknown, limited evidence of the genetics of species' differences suggests that some differences are due to the substitution of single genes of major effect. While neither the genetics of breeds nor the genetics of species provide a perfect model for the genetics of speciation, similarities between the two are sufficiently striking to infer that major, visible, mutations can provide the impetus underlying new directions of sexual selection.

Acoustic Stimulation↗

The sight of the peacock's tail makes me sick: the early arguments on sexual selection.

Why does a peacock have a beautiful train, while a peahen is sober without such flamboyance? Darwin proposed the theory of sexual selection to explain the differences between the sexes of the same species. Recently the study of sexual selection has been one of the most flourishing areas in evolutionary biology. However, the theory met with great resistance from biologists since the publication of the idea and the history of the theory included a lot of misunderstanding and confusion. There are several reasons for this. First, classical Darwinism failed to recognize social competition as an important selective force. Second, the good-for-the-species argument, which persisted in the days after Darwin, made the sexual selection argument more difficult to under-stand. Compared to the discussions on animals, Darwin's argument on human sex differences is not satisfactory. The reason probably lies in the debate over human racial differences which prevailed in the 19th century.

Animals↗

Manifestations of sexual selection may depend on the genetic bases of sex determination.

A variant of the 'handicap' model of sexual selection is described which predicts that the evolution of ornate male traits occurs more easily in species where females are the heterogametic sex. The process occurs even when the alleles conferring high paternal 'fitness' remain advantageous for only a short time due to a rapidly changing physical or biotic environment: the timescale of this advantage may approach the gestation time of the organism. This provides an explanation as to why sexual selection in species where females are heterogametic (such as birds) occurs mainly by the elaboration of ornate male secondary sexual characteristics, whereas in species where females are homogametic (such as mammals) sexual selection results predominantly in inter-male rivalry and the evolution of traits such as horns, antlers and large body size. An analogy between the evolution of elaborate male traits and the evolution of warning coloration is noted.

Alleles↗

Sexual selection and genital anatomy of male primates.

Correlations between mating system and various aspects of genital anatomy suggest a strong influence of sexual selection on genital morphology. We test the generality of the influence by examining whether primate taxa in which there might be enhanced sexual selection (those with multi-male mating systems) possess, as expected, relatively more spinous penises than do taxa with other mating systems. As most prosimians, but few anthropoids (monkeys and apes), possess penile spines, and because the predominant mating systems of the two taxa differ, taxonomic constraints are taken into account. Sexual selection apparently does not act on penile spines in the same manner as on other aspects of genital anatomy: spinosity is not greatest in multi-male taxa of either prosimians or anthropoids. In some taxa, spines might stimulate reproductive readiness and synchrony in situations in which the sexes live apart and do not have other means of communicating reproductive state (dispersed social systems and 'stolen' extra-pair copulations), but problems exist with the hypothesis, as they do with the idea that spines are involved with scent marking. It seems that either penile spines have several functions, or penile spinosity in primates, and other orders, remains to be explained.

Animals↗

Reproductive ageing and sexual selection on male body size in a wild population of antler flies (Protopiophila litigata).

Little is known about the importance of trade-offs between ageing and other life history traits, or the effects of ageing on sexual selection, particularly in wild populations suffering high extrinsic mortality rates. Life history theory suggests that trade-offs between reproduction and somatic maintenance may constrain individuals with higher initial reproductive rates to deteriorate more rapidly, resulting in reduced sexual selection strength. However, this trade-off may be masked by increased condition dependence of reproductive effort in older individuals. We tested for this trade-off in males in a wild population of antler flies (Protopiophila litigata). High mating rate was associated with reduced longevity, as a result of increased short-term mortality risk or accelerated ageing in traits affecting viability. In contrast, large body size was associated with accelerated ageing in traits affecting mating success, resulting in reduced sexual selection for large body size. Thus, ageing can affect sexual selection and evolution in wild populations.

Aging↗

Effectiveness of sexual selection in preventing fitness deterioration in bulb mite populations under relaxed natural selection.

Under the 'good genes' mechanism of sexual selection (SS), females benefit from mate choice indirectly: their offspring inherit genes of the preferred, high quality fathers. Recent models assume that the genetic variance for male quality is maintained by deleterious mutations. Consequently, SS can be predicted to remove deleterious mutations from populations. We tested this prediction by relaxing selection in populations of the bulb mite, thus increasing their rate of accumulation of deleterious mutation. SS, allowed to operate in half of these populations, did not prevent the fitness decline observed in the other half of the relaxed selection lines. After 11 generations of relaxed selection, female fecundity in lines in which males were allowed to compete for females declined compared with control populations by similar amount as in monogamous lines (17.5 and 14.5%, respectively), whereas other fitness components (viability, longevity, male reproductive success) did not differ significantly between both types of lines and control populations.

Analysis of Variance↗