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Rapid evolution of a sexually selected trait following population establishment in a novel habitat.

Colonization of novel environments creates new selection pressures. Sexually selected traits are affected by the physical and social environment and should be especially susceptible to change, but this has rarely been studied. In southern California, dark-eyed juncos, (Junco hyemalis) naturally breed in mixed-coniferous temperate forests, typically from 1500 m to 3000 m in elevation. In the early 1980s, a small population became established in a coastal habitat, the University of California, San Diego campus, which has a mild, Mediterranean climate. I show that a sexually and socially selected signaling trait--the amount of white in the tail--has declined by approximately 22% as compared to mountain juncos. I address three main factors that could explain the difference between mountain and coastal juncos: phenotypic plasticity, genetic drift, and selection. Results indicate that the first two can be ruled out as the sole cause of the plumage change, which implies that selection contributed to the genetic differentiation from the mountain population. The estimated rate of evolution is about 0.2 haldanes, comparable with rates of change in systems where individuals have been artificially introduced into new environments (e.g., guppies and Drosophila). This is the first study to demonstrate evolution of a sexually selected trait after only several generations resulting from a natural invasion into a novel environment.

Animals↗

Parasites, immunology of hosts, and host sexual selection.

Parasite-mediated sexual selection is reviewed with special emphasis on the bird literature. Choosy females may benefit from choosing parasite-free mates if such males provide better parental care, do not transmit contagious parasites, or provide resistance genes to offspring. There is evidence in support of each of these mechanisms. The immunocompetence handicap hypothesis posits that secondary sexual characters reliably reveal the ability of males to resist parasites due to the immunosuppressive effects of testosterone and other biochemicals. Several aspects of these negative feedback mechanisms are supported by laboratory studies, but evidence from free-living animals is almost completely absent. Corticosterone rather than testosterone may potentially mediate the immunocompetence handicap mechanism. A simple version of the immunocompetence handicap is developed suggesting that body condition of male hosts is a sufficient mediator of the handicap mechanism of reliable sexual signaling. Sexual selection appears to be more intense in sexually dichromatic bird species, and comparative studies using pairwise comparisons of closely related taxa reveal that sexually dichromatic bird species have larger spleens, larger bursa of Fabricius, and higher concentrations of leukocytes than monochromatic species. Parasite-mediated sexual selection is proposed to affect parasite biology by increasing (1) the variance-to-mean ratio in parasite abundance, (2) variance in the intensity of natural selection affecting hosts, and (3) speciation rates among parasites exploiting hosts subject to intense sexual selection as compared to those subject to less intense selection.

Animals↗

Reproductive social behavior: cooperative games to replace sexual selection.

Theories about sexual selection can be traced back to Darwin in 1871. He proposed that males fertilize as many females as possible with inexpensive sperm, whereas females, with a limited supply of large eggs, select the genetically highest quality males to endow their offspring with superior capabilities. Since its proposal, problems with this narrative have continued to accumulate, and it is our view that sexual selection theory needs to be replaced. We suggest an approach that relies on the exchange of direct ecological benefits among cooperating animals without reference to genetic benefits. This approach can be expressed mathematically in a branch of game theory that pertains to bargaining and side payments.

Animals↗

Sexual selection for size and symmetry in a diversifying secondary sexual character in Drosophila bipectinata Duda (Diptera: Drosophilidae).

Results of intrapopulation studies of sexual selection and genetic variation and covariation underlying elements of the sex comb of Drosophila bipectinata are presented. The magnitude of the sex comb, a sexual ornament, varies significantly among Australasian populations, motivating research into the evolutionary mechanisms responsible for its incipient diversification. The comb is composed of stout black teeth on the front legs of males arranged in three distinct segments: C1, C2, and C3. Significant sexual selection in field populations in northeastern Queensland, Australia, was detected for increasing C2 and body size, and simultaneously for reducing comb positional fluctuating asymmetry. In contrast, sexual selection was not detected for other comb segments, nor for sternopleural bristle number or symmetry. Selection intensities for C2 and comb positional fluctuating asymmetry were similar in magnitude, and although they were opposite in sign, values across twelve sampling dates, or selection episodes, were uncorrelated. Heritability estimates for C2 were high and significant across years, whereas heritability estimates for comb positional asymmetry were small, and generally nonsignificant. The major sex comb segments (C1 and C2) were significantly and positively correlated genetically, indicating the potential for correlated evolution of these components of the comb under sexual selection. The original finding of a significant positive genetic correlation between the magnitude of this sex trait and its positional asymmetry indicates that the counteracting and independent selection pressures detected could contribute to the maintenance of genetic variation sustaining sexual selection. The study documents the simultaneous presence of sexual selection in nature and of heritable genetic variation underlying expression of the sex comb, fundamental conditions necessary for its adaptive diversification. Drosophila bipectinata may be a valuable model for studies of adaptive diversification and incipient speciation by sexual selection.

Adaptation, Biological↗

Sexual selection in plants: pros and cons.

Sexual selection has long been regarded as a special mode of selection in animals. Various authors have argued in a growing number of publications since 1979 that sexual selection is also operative in plants, and consequently, two divergent concepts of sexual selection are now being stated in the literature, the original Darwinian concept and the new plant-centered concept. An essential feature of the Darwinian concept is the distinction between primary and secondary sexual characters. Sexual selection is a process producing secondary sexual characters in males; such characters are well known in animals but are unknown in plants. Proponents of sexual selection in plants do not mention the subject of primary and secondary sexual characters, and they make no effort to establish the existence of secondary sexual characters in plants. The evidence they do present for sexual selection in plants consists of primary sexual characters and other reproductive traits that are products of selection modes other than sexual selection.

Journal Article↗

Parasitism, host immune function, and sexual selection.

Parasite-mediated sexual selection may arise as a consequence of 1) females avoiding mates with directly transmitted parasites, 2) females choosing less-parasitized males that provide parental care of superior quality, or 3) females choosing males with few parasites in order to obtain genes for parasite resistance in their offspring. Studies of specific host-parasite systems and comparative analyses have revealed both supportive and conflicting evidence for these hypotheses. A meta-analysis of the available evidence revealed a negative relationship between parasite load and the expression of male secondary sexual characters. Experimental studies yielded more strongly negative relationships than observations did, and the relationships were more strongly negative for ectoparasites than for endoparasites. There was no significant difference in the magnitude of the negative effect for species with and without male parental care, or between behavioral and morphological secondary sexual characters. There was a significant difference between studies based on host immune function and those based on parasite loads, with stronger effects for measures of immune function, suggesting that the many negative results from previous analyses of parasite-mediated sexual selection may be explained because relatively benign parasites were studied. The multivariate analyses demonstrating strong effect sizes of immune function in relation to the expression of secondary sexual characters, and for species with male parental care as compared to those without, suggest that parasite resistance may be a general determinant of parasite-mediated sexual selection.

Animals↗

Sexual selection fails to promote adaptation to a new environment.

Selection can be divided into sexual and nonsexual components. Some work finds that a component of sexual selection, adaptive female selection for good genes, can promote nonsexual fitness. Less studied is the benefit from sexual selection in toto, that is, when intra- and intersexual selection are both present and able to affect females directly and indirectly. Here an upper bound for the net benefit of sexual selection is estimated for Drosophila melanogaster. Replicate populations were allowed to adapt to low-grade thermal stress, with or with out the operation of sexual selection. Because proteins and lipids are highly sensitive to temperature, low-grade thermal stress will select broadly across the genome for alternative alleles. Such broad, directional selection for thermal tolerance should increase the measurable benefits of sexual selection far beyond that available under stabilizing selection. Sexual selection was removed by enforced monogamy without mate choice and retained by enforced polyandry (four males per female). After 36 generations of thermal stress exposure, there was substantial adaptation to the new environment (the net reproductive rate increased six standard deviations relative to thermal controls). However, sexual selection did not affect the rate of adaptation. Therefore, adaptive female selection for thermal tolerance either was insignificant or negated by other aspects of sexual selection, for example, male-induced female harm, which has been shown to diminish under monogamy. This experiment employed two parameters that reduced the opportunity for divergence in such harm: a truncated intersexual interaction period and strong directional selection for thermal tolerance. No divergence in male-induced harm was observed.

Adaptation, Physiological↗

Sexual selection: an evolutionary force in plants?

Sexual selection has traditionally been used to explain exaggerated sexual traits in male animals. Today the concept has been developed and various other sexually related traits have been suggested to evolve in the same manner. In nearly all new areas where the theory of sexual selection has been applied, there has been an intense debate as to whether the application is justified. Is it the case that some scientists are all too ready to employ fashionable ideas? Or are there too many dogmatic researchers refusing to accept that science develops and old ideas are transformed? Maybe the controversies are simply a reflection of the difficulty of defining a theory under constant re-evaluation. Thus, we begin by summarizing the theory of sexual selection in order to assess the influence of sexual selection on the evolution of plant morphology. We discuss empirical findings concerning potentially affected traits. Although we have tried to address criticisms fairly, we still conclude that sexual selection can be a useful tool when studying the evolution of reproductive traits in plants. Furthermore, by including the evidence from an additional kingdom, a fuller understanding of the processes involved in sexual selection can be gained.

Biological Evolution↗

Interactions among mechanisms of sexual selection on male body size and head shape in a sexually dimorphic fly.

Darwin envisaged male-male and male-female interactions as mutually supporting mechanisms of sexual selection, in which the best armed males were also the most attractive to females. Although this belief continues to predominate today, it has been challenged by sexual conflict theory, which suggests that divergence in the interests of males and females may result in conflicting sexual selection. This raises the empirical question of how multiple mechanisms of sexual selection interact to shape targeted traits. We investigated sexual selection on male morphology in the sexually dimorphic fly Prochyliza xanthostoma, using indices of male performance in male-male and male-female interactions in laboratory arenas to calculate gradients of direct, linear selection on male body size and an index of head elongation. In male-male combat, the first interaction with a new opponent selected for large body size but reduced head elongation, whereas multiple interactions with the same opponent favored large body size only. In male-female interactions, females preferred males with relatively elongated heads, but male performance of the precopulatory leap favored large body size and, possibly, reduced head elongation. In addition, the amount of sperm transferred (much of which is ingested by females) was an increasing function of both body size and head elongation. Thus, whereas both male-male and male-female interactions favored large male body size, male head shape appeared to be subject to conflicting sexual selection. We argue that conflicting sexual selection may be a common result of divergence in the interests of the sexes.

Aggression↗

The sexually-selected sperm hypothesis: sex-biased inheritance and sexual antagonism.

When females are inseminated by more than one male (polyandry) sexual selection continues after insemination in the form of sperm competition and cryptic female choice. The sexually-selected sperm hypothesis proposes that, under the risk of sperm competition, additive variation in male traits determining fertilising efficiency will select for female propensity to be polyandrous in order to increase the probability of producing sons with superior fertilising efficiency. Two factors complicate this prediction: sex-biased transmission of male fertilising efficiency traits and sexual antagonism of sex-limited traits, fostered by sex-biased inheritance. Here, we (i) review the evidence that male traits contributing towards fertilising efficiency are heritable through sex-biased mechanisms, and (ii) explore the evolutionary implications for male and female reproductive strategies caused by both sex-biased transmission and sexual antagonism of fertilising efficiency traits. Many male fertilising efficiency traits are heritable through sex-biased mechanisms and may not necessarily increase female fitness. The predictions of the sexually-selected sperm hypothesis change dramatically under these different mechanisms of inheritance of fertilising efficiency traits, and different fitness pay-offs derived by females from the expression of such traits. Both sex-biased control of fertilising efficiency and sexual antagonism may also be important in explaining the maintenance of the genetic variance and selection potential of fertilising efficiency. We propose that a useful approach to test the sexually-selected sperm hypothesis is to combine studies which identify behavioural and physiological mechanisms explaining variation in reproductive success with artificial selection experiments to infer the underlying evolutionary patterns.

Animals↗

Faster lizards sire more offspring: sexual selection on whole-animal performance.

Sexual selection operates by acting on variation in mating success. However, since selection acts on whole-organism manifestations (i.e., performance) of underlying morphological traits, tests for phenotypic effects of sexual selection should consider whole-animal performance as a substrate for sexual selection. Previous studies have revealed positive relationships between performance and survival, that is, natural selection, but none have explicitly tested whether performance may influence reproductive success (through more matings), that is, sexual selection. Performance predicts dominance in some species, implying the effects of sexual selection, but how it does so has not been established, nor is it certain whether performance might be a by-product of selection for something else, for example, elevated circulating testosterone levels. We investigated the potential for sexual selection on sprint speed performance in collared lizards (Crotaphytus collaris), considering the potential mediating effects of circulating hormone levels. Among territorial, adult male collared lizards, only sprint speed significantly predicted territory area and number of offspring sired as determined by genetic paternity analysis. Body size, head size, and hind limb length had no effect. Neither plasma testosterone levels nor corticosterone levels correlated with sprint speed, territory area, or number of offspring sired. Thus, our results provide a direct link between whole-animal performance and reproductive success, suggesting that intrasexual selection can act directly on sprint speed performance and drive the evolution of underlying morphological traits.

Animals↗

Signal trait sexual dimorphism and mutual sexual selection in Drosophila serrata.

The evolution of sexual dimorphism may occur when natural and sexual selection result in different optimum trait values for males and females. Perhaps the most prominent examples of sexual dimorphism occur in sexually selected traits, for which males usually display exaggerated trait levels, while females may show reduced expression of the trait. In some species, females also exhibit secondary sexual traits that may either be a consequence of a correlated response to sexual selection on males or direct sexual selection for female secondary sexual traits. In this experiment, we simultaneously measure the intersex genetic correlations and the relative strength of sexual selection on males and females for a set of cuticular hydrocarbons in Drosophila serrata. There was significant directional sexual selection on both male and female cuticular hydrocarbons: the strength of sexual selection did not differ among the sexes but males and females preferred different cuticular hydrocarbons. In contrast with many previous studies of sexual dimorphism, intersex genetic correlations were low. The evolution of sexual dimorphism in D. serrata appears to have been achieved by sex-limited expression of traits controlled by genes on the X chromosome and is likely to be in its final stages.

Animals↗

Quantitative genetic models of sexual selection: a review.

Quantitative genetic models of sexual selection have disproven some of the central tenets of both the handicap mechanism and the 'sexy son' hypothesis. These results suggest that the 'good genes' approach to sexual selection may often lead to erroneous results. Runaway sexual selection seems possible under a wide variety of circumstances. Quantitative genetic models have revealed runaway processes for sexually selected attributes expressed in both sexes and for attributes of parental care. Furthermore, the runaway could occur simultaneously in a series of populations that straddle an environmental gradient. While the models support the feasibility of runaway processes, empirical studies are needed to evaluate whether runaways actually happen. Estimates of critical genetic parameters are particularly needed, as well as measures of natural and sexual selection acting on the same population. The models also show that sexual selection has tremendous potential to produce population differentiation, particularly in epigamic traits. Differentiation is promoted by indeterminancy of evolutionary outcome, transient differences among populations during the final slow approach to equilibrium, sampling drift among equilibrium populations, and the tendency of sexual selection to amplify geographic variation arising from spatial differences in natural selection. Recent work with two- and three-locus models of sexual selection has produced results that parallel the results of the polygenic models (Kirkpatrick, 1982, 1985, 1986; Seger, 1985). Thus the feature of indeterminate equilibria (outcome dependent on initial conditions) is common to both types of model.

Animals↗

Human skin-color sexual dimorphism: a test of the sexual selection hypothesis.

Applied to skin color, the sexual selection hypothesis proposes that male preference for light-skinned females explains the presence of light skin in areas of low solar radiation. According to this proposal, in areas of high solar radiation, natural selection for dark skin overrides the universal preference of males for light females. But in areas in which natural selection ceases to act, sexual selection becomes more important, and causes human populations to become light-skinned, and females to be lighter than males. The sexual selection hypothesis proposes that human sexual dimorphism of skin color should be positively correlated with distance from the equator. We tested the prediction that sexual dimorphism should increase with increasing latitude, using adult-only data sets derived from measurements with standard reflectance spectrophotometric devices. Our analysis failed to support the prediction of a positive correlation between increasing distance from the equator and increased sexual dimorphism. We found no evidence in support of the sexual selection hypothesis.

Female↗

Broad-sense sexual selection, sex gene pool evolution, and speciation.

Studies of sexual selection have traditionally focused on explaining the extreme sexual dimorphism in male secondary sexual traits and elaborate mating behaviors displayed by males during courtship. In recent years, two aspects of sexual selection have received considerable attention in the literature: an extension of the sexual selection concept to other traits (i.e., postcopulatory behaviors, external and internal genital morphology, gametes, molecules), and alternative mechanistic explanations of the sexual selection process (i.e., coevolutionary runaway, good-genes, sexual conflicts). This article focuses on the need for an extension of sexual selection as a mechanism of change for courtship and (or) mating male characters (i.e., narrow-sense sexual selection) to all components of sexuality not necessarily related to courtship or mating (i.e., broad-sense sexual selection). We bring together evidence from a wide variety of organisms to show that sex-related genes evolve at a fast rate, and discuss the potential role of broad-sense sexual selection as an alternative to models that limit speciation to strict demographic conditions or treat it simply as an epiphenomenon of adaptive evolution.

Animals↗

Sexually selected traits and adult survival: a meta-analysis.

Traits correlated with male mating success are likely to be subject to sexual selection. Sexually selected characters are thought to be costly to develop and maintain. If males do not vary their investment in sexual traits in relation to their ability to bear the costs, there should be a negative relationship between male longevity or survival and the expression of sexual traits. In particular, a negative relationship is predicted by pure Fisherian models for the evolution of sexual ornaments. The same should also be true for traits that evolve via pleiotropy (e.g., due to sensory exploitation or bias) with no subsequent evolution of condition dependent modification. We collected information on the relationship between traits correlated with male mating rate and estimates of adult male survivorship or life span. In total we obtained 122 samples from 69 studies of 40 species of bird, spider, insect, and fish. In a meta-analysis we calculated the average sample size weighted correlation between trait expression and adult survival. Analyses at the level of samples, studies, and species revealed significant positive relationships (r = 0.08, 0.10, and 0.13, respectively; all P < 0.001). The unweighted correlation at the species level was r = 0.24. In general, males with larger ornaments or weapons, greater body size, or higher rates of courtship showed greater survivorship or longevity. This finding is inconsistent with pure Fisherian models or other models that do not incorporate condition or quality dependent trait expression. It suggests that male investment in sexually selected traits is not fixed but varies in relation to the ability to pay the underlying costs of expressing these characters. Hence, many secondary sexual characters are likely to be condition dependent in their expression.

Animals↗

Sexual selection and the risk of extinction in mammals.

Sexual selection is commonly envisaged as a force working in opposition to natural selection, because extravagant or exaggerated traits could apparently have never evolved via natural selection alone. There is good evidence that a selection load imposed by sexual selection may be eased experimentally by restricting the opportunity for it to operate. Sexual selection could therefore potentially play an important role in influencing the risk of extinction that a population faces, thereby contributing to the apparent selectivity of extinctions. Conversely, recent theory predicts that the likelihood of extinction may decrease when sexual selection is operating because it could accelerate the rate of adaptation in concert with natural selection. So far, comparative evidence (coming mostly from birds) has generally indicated support for the former scenario, but the question remains open. The aim of this study was therefore to examine whether the level of sexual selection (measured as residual testes mass and sexual size dimorphism) was related to the risk of extinction that mammals are currently experiencing. We found no evidence for a relationship between these factors, although our analyses may have been confounded by the possible dominating effect of contemporary anthropogenic factors.

Animals↗

Sexual selection and the risk of extinction in birds.

The relationship between sexual selection and extinction risk has rarely been investigated. This is unfortunate because extinction plays a key role in determining the patterns of species richness seen in extant clades, which form the basis of comparative studies into the role that sexual selection may play in promoting speciation. We investigate the extent to which the perceived risk of extinction relates to four different estimates of sexual selection in 1030 species of birds. We find no evidence that the number of threatened species is distributed unevenly according to a social mating system, and neither of our two measures of pre-mating sexual selection (sexual dimorphism and dichromatism) was related to extinction risk, after controlling for phylogenetic inertia. However, threatened species apparently experience more intense post-mating sexual selection, measured as testis size, than non-threatened species. These results persisted after including body size as a covariate in the analysis, and became even stronger after controlling for clutch size (two known correlates of extinction risk). Sexual selection may therefore be a double-edged process-promoting speciation on one hand but promoting extinction on the other. Furthermore, we suggest that it is post-mating sexual selection, in particular, that is responsible for the negative effect of sexual selection on clade size. Why this might be is unclear, but the mean population fitness of species with high intensities of post-mating sexual selection may be especially low if costs associated with multiple mating are high or if the selection load imposed by post-mating selection is higher relative to that of pre-mating sexual selection.

Animals↗