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Interspecific aggression causes negative selection on sexual characters.

Interspecific aggression originating from mistaken species recognition may cause selection on secondary sexual characters, but this hypothesis has remained untested. Here we report a field experiment designed to test directly whether interspecific aggression causes selection on secondary sexual characters, wing spots, in wild damselfly populations. Males of Calopteryx virgo are more aggressive toward males of C. splendens with large than with small wing spots. This differential interspecific aggression may cause negative selection on wing spot size. Indeed, our results show that directional survival selection on wing spot size of C. splendens males was changed by experimental removal of C. virgo males. Without removal, directional selection went from positive to negative with increasing relative abundance of C. virgo males. In populations where C. virgo males were removed, this relationship disappeared. These results verify that interspecific aggression can cause negative selection on sexual characters. Thus, interspecific aggression has the potential to cause divergence on these characters between two species offering an alternative explanation for reinforcement for generating character displacement in secondary sexual characters.

Aggression↗

Sexual partner selectiveness effects on homosexual HIV transmission dynamics.

Deterministic simulation models are used to show that HIV transmission dynamics in homosexual populations can be strongly affected by sexual partner selectiveness. The type of selectiveness or biased mixing examined is where individuals with similar new partnership formation rates are more likely to form a pair than would be expected by chance. The effect of such selectiveness could be strong even when the total number and distribution of new sexual partnerships and sex acts remains constant. This means that in order to predict the future course of HIV transmission and identify the populations at highest risk, we must have information not only on the frequency of new sexual partnerships and types of sex acts, but also on who has sex with whom. Given high sexual partner selectiveness, some groups of homosexuals with low rates of sex and new sex partners would take many decades before a single introduction would generate an epidemic. Epidemics in these groups can be markedly accelerated by only modest contact with higher risk groups. Even in very low activity groups, which if isolated would have no epidemic, an important proportion of their members can be infected when they are not selective. The relative risks of AIDS in groups making high numbers of new sexual partnerships compared to groups making low numbers are markedly affected by sexual partner selectiveness. The models developed were examined using information collected in 1984 from the Coping and Change Study in collaboration with the Chicago Multicenter AIDS Cohort Study. This population was divided into activity groups by the rate at which individuals established new sexual partnerships and then a structure of new sexual partnerships between these activity groups was defined consistent with available data. Even without introducing any behavior change in the models, the proportion of the homosexual population infected was seen to level off temporarily at around 50% after several years as a consequence of saturation in the high risk groups when lower risk groups were not yet being consumed by the epidemic process. Most sex in the study population is casual or anonymous. The sample selection procedures were biased toward individuals who engage in such sex. There is, however, a large group of individuals that avoids the casual and anonymous sex scene. Simulated epidemics indicated that under a variety of conditions, the population that does not engage in casual and anonymous sex could experience an epidemic of HIV infection quite some time after the epidemic in the population that does has waned.(ABSTRACT TRUNCATED AT 400 WORDS)

Acquired Immunodeficiency Syndrome↗

Sexual responsiveness is condition-dependent in female guppies, but preference functions are not.

BACKGROUND: Variation in mate choice behaviour among females within a population may influence the strength and form of sexual selection, yet the basis for any such variation is still poorly understood. Condition-dependence may be an important source of variation in female sexual responsiveness and in the preference functions for male display traits that she expresses when choosing. We manipulated food intake of female guppies (Poecilia reticulata), and examined the effect on several measures of condition and various components of mate choice behaviour. RESULTS: Diet significantly influenced four measures of female condition: standard length, weight, reproductive status and somatic fat reserves. Diet also significantly affected female sexual responsiveness, but not preference functions: females in good and poor condition prefer the same males. CONCLUSIONS: Variation in female condition within populations is therefore unlikely to influence the direction of sexual selection imposed by female choice. It may, however, influence the strength of sexual selection due to its effects on female responsiveness. The relative importance of female choice as a sexually selective force may also covary with female condition, however, because low responsiveness may result in sneak copulations being relatively more important as a determinant of the paternity of offspring. Differences among populations in mean condition may also influence geographic differences in the strength of sexual selection.

Analysis of Variance↗

Molecular parentage analysis in experimental newt populations: the response of mating system measures to variation in the operational sex ratio.

Molecular studies of parentage have been extremely influential in the study of sexual selection in the last decade, but a consensus statistical method for the characterization of genetic mating systems has not yet emerged. Here we study the utility of alternative mating system measures by experimentally altering the intensity of sexual selection in laboratory-based breeding populations of the rough-skinned newt. Our experiment involved skewed sex ratio (high sexual selection) and even sex ratio (low sexual selection) treatments, and we assessed the mating system by assigning parentage with microsatellite markers. Our results show that mating system measures based on Bateman's principles accurately reflect the intensity of sexual selection. One key component of this way of quantifying mating systems is the Bateman gradient, which is currently underutilized in the study of genetic mating systems. We also compare inferences based on Bateman's principles with those obtained using two other mating system measures that have been advocated recently (Morisita's index and the index of resource monopolization), and our results produce no justification for the use of these alternative measures. Overall, our results show that Bateman's principles provide the best available method for the statistical characterization of mating systems in nature.

Animals↗

Components of lifetime mating success and body size in males of a scrambling damselfly.

Sexual selection is hypothesized to favour small body size in males of scrambling species, that is, those in which males obtain matings by actively searching for females. I tested this hypothesis in a natural population of the scrambling emerald damselfly, Lestes sponsa. Mating efficiency (matings/visit to the pond) was the most important factor explaining variation in male lifetime mating success (LMS; 71%). This suggests a large potential for sexual selection. Path analysis of male LMS suggested a quality factor that positively affected both mating efficiency and life span. In contrast with the small-male mating advantage hypothesis, part of this potential for sexual selection was realized as stabilizing selection on male body size, indicating that there may also be a lower limit to body size for mating efficiency. This also illustrates that the constancy of body size may be explained by sexual selection alone. Survival explained about 20% of the variation in LMS and random processes were potentially important for determining LMS. My results show the problems of using mating efficiency as a measure for the intensity of sexual selection and the need to distinguish between potential and realized selection pressures, especially when comparing the importance of natural and sexual selection. I discuss mechanisms that may have caused the intermediate-male mating advantage in this scrambling species. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Mental traits as fitness indicators. Expanding evolutionary psychology's adaptationism.

According to most evolutionary psychologists, human psychological adaptations can be recognized by criteria such as high efficiency, high complexity, high modularity, low phenotypic variance, low genotypic variance, low heritability, universality across cultures, and universality across individuals. These criteria are appropriate for adaptations that have been shaped through stabilizing selection for survival utility. However, they are often inappropriate for adaptations that have been shaped by sexual selection through mate choice as reliable signals of heritable fitness. If some psychological adaptations evolved as sexually selected fitness indicators of this type, we should expect them to violate many standard criteria used by evolutionary psychology to distinguish adaptations from non-adaptations. This paper addresses the problems raised by new developments in sexual selection theory and animal signaling theory for evolutionary psychology's adaptationism. It suggests that our adaptationist criteria must recognize two typical kinds of psychological adaptations: naturally selected survival mechanisms and sexually selected fitness indicators.

Adaptation, Psychological↗

Melanin-based plumage coloration and flight displays in plovers and allies.

Plovers and their allies exhibit an impressive diversity of melanin-based plumage patterns ranging from non-melanized to completely melanized species. We use phylogenetic comparative methods to test whether melanization has evolved in relation to sexual selection for attracting mates, to selection for signalling territory defence, or to natural selection for camouflage. First, according to sexual-selection theory, melanized plumage has evolved to amplify the courtship displays of males. As predicted by this hypothesis, we found that males with aerial displays had more melanized plumage than males of ground-displaying species. In addition, sexual dimorphism in melanization was greater in species with display flights than in species with ground displays. Second, melanization may have evolved through social interactions to signal competitive ability in territory defence. We did not find evidence for this hypothesis, since breeding density was unrelated to the melanization of either sex. Finally, melanized plumage may camouflage the incubating parent. The latter hypothesis was not supported, since melanization was unrelated either to the darkness of nest substrate or the extent of vegetation cover. Taken together, our results are most consistent with the sexual-selection hypothesis, and suggest that melanized plumage has evolved to enhance the aerial displays of male plovers.

Animal Communication↗

Song variation in an avian ring species.

Divergence of mating signals can occur rapidly and be of prime importance in causing reproductive isolation and speciation. A ring species, in which two reproductively isolated taxa are connected by a chain of intergrading populations, provides a rare opportunity to use spatial variation to reconstruct the history of divergence. I use geographic variation in the song of a likely ring species, the greenish warbler (Phylloscopus trochiloides) to reconstruct the microevolutionary steps that occurred during divergence of a trait that is often important in speciation in birds. Populations of a western Siberian (P. t. viridanus) and an eastern Siberian (P. t. plumbeitarsus) form of the greenish warbler meet, but do not interbreed in central Siberia; these forms are connected by a chain of interbreeding populations extending in a ring to the south around the treeless Tibetan Plateau. I show that: (1) song structure differs greatly between the two Siberian forms, which share the same habitat; (2) song structure changes gradually around the ring; (3) singing behavior is relatively simple in the Himalayas, but becomes increasingly complex to the north, both to the west and east of the Tibetan Plateau; and (4) song varies along independent axes of complexity in the western and eastern south-north clines. By comparing geographic variation in singing behavior and ecological variables, I distinguish among possible causes of song divergence, including selection based on the acoustic environment, stochastic effects of sexual selection, and selection for species recognition. I suggest that parallel south-to-north ecological gradients have caused a greater intensity of sexual selection on song in northern populations and that the stochastic effects of sexual selection have led to divergence in song structure.

Animals↗

Male genital size reflects a tradeoff between attracting mates and avoiding predators in two live-bearing fish species.

Male genitalia may experience more rapid, divergent evolution than any other animal character, but why? Research during the past several decades has culminated in the view that genital diversification primarily results from postmating sexual selection (e.g., sperm competition or cryptic female choice). However, the potential roles of premating sexual selection (e.g., mate choice) and natural selection have received little attention. We examined the possible importance of these mechanisms by investigating divergence in male genitalia among populations differing in predator regime for two species of live-bearing fish (Gambusia affinis in Texas and Gambusia hubbsi in The Bahamas). When controlled for body size, males exhibited a larger gonopodium (sperm-transfer organ) in predator-free environments than in predatory environments, a trend that persisted across space (multiple populations), time (multiple years), and species. By conducting laboratory experiments with G. affinis, we found that premating sexual selection seems to favor larger male genitalia (females exhibited mating preference for males having larger gonopodia), but natural selection in the presence of predatory fishes seems to favor reduced genital size (larger gonopodium size was associated with reduced burst-swimming performance, an important antipredator behavior). Although postmating sexual selection is widely presumed to be the most important mechanism driving genital diversification, these findings suggest that alternative mechanisms, particularly for organisms that cannot retract their genitalia, may also prove important.

Analysis of Variance↗

Relative fitness can decrease in evolving asexual populations of S. cerevisiae.

It is generally accepted from the darwinian theory of evolution that a progressive increase in population adaptation will occur in populations containing genetic variation in fitness, until a stable equilibrium is reached and/or the additive genetic variation is exhausted. However, the theoretical literature of population genetics documents exceptions where mean population fitness may decrease in response to evolutionary changes in gene frequency, due to varying selective coefficients, sexual selection or to epistatic interactions between loci. Until now, no examples of such exceptions have been documented from fitness estimates in either natural or experimental populations. We present here direct evidence that, as a result of epistatic interactions between adaptive mutations, mean population fitness can decrease in asexual evolving populations of the yeast Saccharomyces cerevisiae.

Biological Evolution↗

Conspecific sperm precedence in Drosophila.

Traits that influence the interactions between males and females can evolve very rapidly through sexual selection or sexually antagonistic coevolution. Rapid change in the fertilization systems of independent populations can give rise to reproductive incompatibilities between populations, and may contribute to speciation. Here I provide evidence for cryptic reproductive divergence among three sibling species of Drosophila that leads to a form of postmating isolation. When a female mates with both a conspecific and a heterospecific male, the conspecific sperm fertilize the vast majority of the eggs, regardless of the order of the matings. Heterospecific sperm fertilize fewer eggs after these double matings than after single matings. Experiments using spermless males show that the seminal fluid of the conspecific male is largely responsible for this conspecific sperm precedence. Moreover, when two males of the same species mate sequentially with a female from a different species, a highly variable pattern of sperm precedence replaces the second-male sperm precedence that is consistently found within species. These results indicate that females mediate sperm competition, and that second-male sperm precedence is not an automatic consequence of the mechanics of sperm storage.

Animals↗

Human vision fails to distinguish widespread sexual dichromatism among sexually "monochromatic" birds.

Historical scenarios of evolution of avian plumage coloration have been called into question with the discoveries that most birds can see UV light (which normal humans cannot), and that UV-reflecting plumages are widespread in birds. Several examples of sexual dichromatism not detectable with human visual capabilities suggest that our categorizations of plumages as sexually mono- or dichromatic might often be incorrect. Nonetheless, given the limited taxonomic scope of those examples, the vast majority of sexually monochromatic birds are still treated as such without question in avian research. Herein, I show that >90% of 139 species, in a broad sampling of presumed sexually monochromatic passerine birds, were actually sexually dichromatic from an avian visual perspective, based on comparisons of plumage reflectance data using a visual model of color discrimination thresholds. The taxonomic ubiquity of this result suggests that many existing interpretations of evolutionary patterns of sexual dichromatism in birds are erroneous. The visual model used herein provides a method for quantifying sexual dichromatism, revealing that most (58.7%) feather patches sampled lie along a continuum of dichromatism between avian and human discriminatory abilities and could represent unrecognized sexually selected signals. Sexual dichromatism in this study rarely resulted from intersexual differences in UV coloration alone, emphasizing the need for analysis of bird coloration in relation to the full extent of avian visual discriminatory abilities, including, but not limited to, UV-visual capabilities.

Animals↗

The evolution of body size: what keeps organisms small?

It is widely agreed that fecundity selection and sexual selection are the major evolutionary forces that select for larger body size in most organisms. The general, equilibrium view is that selection for large body size is eventually counterbalanced by opposing selective forces. While the evidence for selection favoring larger body size is overwhelming, counterbalancing selection favoring small body size is often masked by the good condition of the larger organism and is therefore less obvious. The suggested costs of large size are: (1) viability costs in juveniles due to long development and/or fast growth; (2) viability costs in adults and juveniles due to predation, parasitism, or starvation because of reduced agility, increased detectability, higher energy requirements, heat stress, and/or intrinsic costs of reproduction; (3) decreased mating success of large males due to reduced agility and/or high energy requirements; and (4) decreased reproductive success of large females and males due to late reproduction. A review of the literature indicates a substantial lack of empirical evidence for these various mechanisms and highlights the need for experimental studies that specifically address the fitness costs of being large at the ecological, physiological, and genetic levels. Specifically, theoretical investigations and comprehensive case studies of particular model species are needed to elucidate whether sporadic selection in time and space is sufficient to counterbalance perpetual and strong selection for large body size.

Adaptation, Physiological↗

Reduced sequence variability on the Neo-Y chromosome of Drosophila americana americana.

Sex chromosomes are generally morphologically and functionally distinct, but the evolutionary forces that cause this differentiation are poorly understood. Drosophila americana americana was used in this study to examine one aspect of sex chromosome evolution, the degeneration of nonrecombining Y chromosomes. The primary X chromosome of D. a. americana is fused with a chromosomal element that was ancestrally an autosome, causing this homologous chromosomal pair to segregate with the sex chromosomes. Sequence variation at the Alcohol Dehydrogenase (Adh) gene was used to determine the pattern of nucleotide variation on the neo-sex chromosomes in natural populations. Sequences of Adh were obtained for neo-X and neo-Y chromosomes of D. a. americana, and for Adh of D. a. texana, in which it is autosomal. No significant sequence differentiation is present between the neo-X and neo-Y chromosomes of D. a. americana or the autosomes of D. a. texana. There is a significantly lower level of sequence diversity on the neo-Y chromosome relative to the neo-X in D. a. americana. This reduction in variability on the neo-Y does not appear to have resulted from a selective sweep. Coalescent simulations of the evolutionary transition of an autosome into a Y chromosome indicate there may be a low level of recombination between the neo-X and neo-Y alleles of Adh and that the effective population size of this chromosome may have been reduced below the expected value of 25% of the autosomal effective size, possibly because of the effects of background selection or sexual selection.

Alcohol Dehydrogenase↗

Number of sexual partners, condom use and risk of human immunodeficiency virus infection.

BACKGROUND: To analyse the relation between number of sexual partners, selected sexual habits and the risk of human immunodeficiency virus (HIV) infection. METHODS: We conducted a case-control study nested in a cross-sectional survey conducted among subjects attending sexually transmitted diseases (STD) clinics in Northern Italy. Eligible for the study were 1711 subjects (1259 males, 452 females) who referred themselves for the first time between September 1988 and March 1993 to three STD clinics in Northern Italy for suspected STD or STD treatment. A total of 145 subjects (113 males and 32 females) were HIV positive. RESULTS: In comparison with subjects reporting no or one sexual partner over the 3 years before the interview, the estimated odds ratios (OR) of HIV serum positivity were 1.2 (95% confidence interval [CI]: 0.6-2.3), 0.8 (95% CI: 0.4-1.8) and 0.3 (95% CI: 0.4-2.5) in subjects reporting 2-3, 4-5, and > or = 6 partners, respectively. The results were similar considering separately males and females and in men reporting only homosexual partners. Regular condom use decreased the risk of HIV infection: in comparison with subjects reporting no or occasional use of condoms, the OR of HIV infection was 0.5 (95% CI: 0.4-0.8) for regular users. Considering men only, compared with men with no homosexual intercourse, the OR of HIV infection was 2.3 (95% CI: 1.4-3.9) in those reporting bisexual intercourse and 2.2 (95% CI: 1.2-4.2) in men reporting only homosexual intercourse (among homosexuals). There was no relation between HIV infection risk and receptive anal sex. CONCLUSIONS: The risk of HIV infection does not increase linearly with the number of sexual partners in this population. This is reasonable, as the prevalence of HIV infection in this population is essentially determined by drug use. Caution is needed in the interpretation of these results since the analysis of role of number of sexual partners in male intravenous drug users is impaired by low statistical power.

Adult↗

Pectoral fins and paternal quality in sticklebacks.

Sexual selection through female mate choice exerts a strong selection pressure on males' sexual traits, particularly when direct benefits are involved. In species with male parental care, one would expect sexual selection to favour paternal quality, for instance through selection on morphological structures which promote quality. We experimentally studied the influence of pectoral fins on paternal quality in male three-spined sticklebacks (Gasterosteus aculeatus L.). After reductions of fin area to different degrees, similar-sized males had to perform a complete reproductive cycle in enclosures in the field. The collected data on fanning behaviour and egg development showed that a reduction in pectoral fin size affected paternal quality probably through an increased beat frequency of the pectorals. Thus, pectoral fins can potentially signal paternal quality to choosy females.

Animals↗

The relationship between conspecific fertilization success and reproductive isolation among three congeneric sea urchins.

Few data are available on the effectiveness of reproductive isolating mechanisms in externally fertilizing taxa. I investigated patterns of conspecific and heterospecific fertilization among three coexisting sea urchin species, Strongylocentrotus droebachiensis, S.franciscanus, and S. purpuratus. In the laboratory, both among and within species, eggs from individual females whose eggs are more easily fertilized by conspecific sperm are also most susceptible to heterospecific fertilization. At one extreme, S. droebachiensis requires an order of magnitude fewer conspecific sperm to fertilize eggs than do the other two species and shows very little distinction between conspecific and heterospecific sperm in no choice experiments. Strongylocentrotus franciscanus has an intermediate susceptibility to fertilization by heterospecific sperm. At the other extreme, S. purpuratus rarely cross-fertilizes. Field observations indicate that S. droebachiensis is often surrounded by heterospecific sea urchins. Genetic analysis of larvae produced during heterospecific spawning events indicate that hybrids are generally produced if male conspecifics are more than 1 m from a spawning female S. droebachiensis. Laboratory cultures indicate that these hybrids suffer high mortality relative to conspecific larvae. Comparisons of reproductive success of S. droebachiensis during single-species and multispecies spawning events indicate that the benefits of producing easily fertilized eggs under conditions of sperm limitation may outweigh the costs of losing some offspring to hybrid fertilization. Patterns of variability in heterospecific fertilization are considered in light of three hypotheses: phylogenetic relatedness, reinforcement selection, and sexual selection.

Animals↗

Measuring polyandry in wild populations: a case study using promiscuous crickets.

Mating rate has important implications for patterns of sexual selection and sexual conflict and hence for issues such as speciation and the maintenance of genetic diversity. Knowledge of natural mating rates can provide insights into the factors driving female mating behaviour. We investigated the level of polyandry in a Spanish population of the field cricket Gryllus bimaculatus using microsatellite markers. Two approaches were employed: (i) genotyping the offspring of wild-caught gravid females to determine the number of males siring the brood and (ii) genotyping sperm stored in the spermathecae of females mated in the wild to estimate the number of mating partners. We compared existing methods for inferring the minimum and probable number of fathers and described a novel probabilistic technique estimating the number of mates by genotyping stored sperm. Using the most conservative allele-counting method, 71% of females produced offspring sired by at least two males (a minimum mean of 2.4 fathers per clutch), and all females had mated to at least two males with minimum mean estimates of 2.7-5.1 mates per female. Our study reveals high levels of polyandry in the wild and suggests that females mate with more males than sire their offspring.

Animals↗