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Sexual size dimorphism in shorebirds, gulls, and alcids: the influence of sexual and natural selection.

Charadrii (shorebirds, gulls, and alcids) have an unusual diversity in their sexual size dimorphism, ranging from monomorphism to either male-biased or female-biased dimorphism. We use comparative analyses to investigate whether this variation relates to sexual selection through competition for mates or natural selection through different use of resources by males and females. As predicted by sexual selection theory, we found that in taxa with socially polygynous mating systems, males were relatively larger than females compared with less polygynous species. Furthermore, evolution toward socially polyandrous mating systems was correlated with decreases in relative male size. These patterns depend on the kinds of courtship displays performed by males. In taxa with acrobatic flight displays, males are relatively smaller than in taxa in which courtship involves simple flights or displays from the ground. This result remains significant when the relationship with mating system is controlled statistically, thereby explaining the enigma of why males are often smaller than females in socially monogamous species. We did not find evidence that evolutionary changes in sexual dimorphism relate to niche division on the breeding grounds. In particular, biparental species did not have greater dimorphism in bill lengths than uniparental species, contrary to the hypothesis that selection for ecological divergence on the breeding grounds has been important as a general explanation for patterns of bill dimorphism. Taken together, these results strongly suggest that sexual selection has had a major influence on sexual size dimorphism in Charadrii, whereas divergence in the use of feeding resources while breeding was not supported by our analyses.

Animals↗

Intrasexual competition and canine dimorphism in anthropoid primates.

A number of factors, including sexual selection, body weight, body-weight dimorphism, predation, diet, and phylogenetic inertia have been proposed as influences on the evolution of canine dimorphism in anthropoid primates. Although these factors are not mutually exclusive, opinions vary as to which is the most important. The role of sexual selection has been questioned because mating system, which should reflect its strength, poorly predicts variation in canine dimorphism, particularly among polygynous species. Kay et al. (1988) demonstrate that a more refined estimate of intermale competition explains a large proportion of the variation in canine dimorphism in platyrrhine primates. We expand their analysis, developing a more generalized measure of intermale competition based on the frequency and intensity of male-male agonism. We examine the relative influences of predation (inferred by substrate use), female body weight, body-weight dimorphism, diet, and sexual selection on the evolution of anthropoid canine dimorphism. Intermale competition is very strongly associated with canine dimorphism. Predation also has a marked effect on canine dimorphism, in that savanna-dwelling species consistently show greater canine dimorphism than other species, all other factors being held equal. Body-weight dimorphism is also strongly associated with canine dimorphism, though apparently through a common selective basis, rather than through allometric effects. Body weight seems to play only a minor, indirect role in the evolution of canine dimorphism. Diet plays no role. Likewise, we find little evidence that phylogenetic inertia is a constraint on the evolution of canine dimorphism.

Analysis of Variance↗

Models of selective mating and the initiation of the Fisherian process.

The effects of various rules of selective mating on the initial stages of Fisherian sexual selection are investigated. A comparison of three models of selective mating, fixed relative preference, best of N males and absolute preference is provided, with a special emphasis on their mathematical properties. Using a two-locus haploid model of sexual selection in a polygamous population, I show that the absolute preference rule of selective mating may lower the threshold frequency of the preference trait, required for the initiation of the Fisherian process, as low as zero. This was not observed in the previous analyses with fixed relative preference or best of N male rules. It is then argued that absolute preference may cause the initiations of the Fisherian process more easily without introducing additional assumptions such as pleiotropy or random genetic drift. Some problems associated with the mating rule are also discussed.

Animals↗

Sexual signaling and speciation, a microevolutionary perspective.

Despite the growing evidence that sexual selection can drive speciation, the evolution of sexual signals in natural populations is far from being well-understood. Sexual signals evolve in response to a variety of factors. Some of the most important selective factors are conspecifics, transmission efficiency in a particular environment, detection by predators, and phylogenetic constraints. These factors have been addressed quite successfully in studies of single types of signals in both vertebrates and invertebrates. However, it is less clear how multimodal signals evolve because the factors listed above will act on every component of the signaling system, and the relative weights of each type of signal must be taken into account. Species of Drosophila are excellent for such analyses because they are amenable to both phenotypic experimentation and genetic manipulation. This paper presents an approach that involves two analyses: studies of which signals are sexually selected within a species, and parallel studies of the signals that are involved in behavioral isolation between closely related species. If the same signal characteristics are involved in both processes, they would provide support for the hypothesis that sexual selection can drive speciation. This approach is illustrated with studies of Hawaiian Drosophila and a review of signals that could be sexually selected in Drosophila melanogaster.

Animals↗

A test of sensory bias for long tails in the house finch.

The sensory bias model of sexual selection proposes that female preferences for certain male traits result from biases in the female sensory system that existed prior to the evolution of the preferred trait. Phylogenetic tests of the sensory bias hypothesis have provided evidence for pre-existing bias in several species, but the hypothesis has not been tested relative to elongated tail feathers, a widespread sexually selected trait in birds. Elongated tails occur sporadically across avian taxa, including cardueline finches (family Fringillidae), suggesting that long tails evolved independently in many lineages. To determine whether a female preference for longer tails predated the evolution of long tails in cardueline finches, we tested female preferences for artificially lengthened tail feathers in male house finches, Carpodacus mexicanus. The house finch is a cardueline finch species with a tail that does not appear to be elongated or sexually selected. In mate choice experiments, females did not demonstrate a preference for lengthened tails over controls. This negative result fails to support the hypothesis that elongation of tail feathers resulted from exploitation of a widespread sensory bias. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Functional significance of phytochemical lures to dacine fruit flies (Diptera: Tephritidae): an ecological and evolutionary synthesis.

While lures of plant origin are vital tools in dacine (Diptera: Tephritidae) pest management, the ecological and evolutionary significance of this lure response remains enigmatic. Two hypotheses (the ancestral host hypothesis and sexual selection by female choice) have been invoked to explain the functional significance of these chemicals to dacine fruit flies. These hypotheses are often treated as alternatives to one another and evidence favouring one is used to reject the other. This review highlights that these two hypotheses are not logical alternatives to each other as the ancestral host hypothesis attempts to explain the ultimate function of the response of Dacinae to these plant-derived parapheromones while the sexual selection hypothesis provides a proximate explanation for lure response. Research on lure response, dacine mating behaviour, functional significance of lures, plant phylogeny and biochemistry and dacine pheromone chemistry are used to evaluate the evidence in relation to both these hypotheses. Some of the key findings are that there is evidence both in support of and against these two hypotheses. Response of fruit flies to related phenyl propanoids to those commonly used as lures in pest management and distribution of phenyl propanoids attractive to dacines among plant orders strongly support the ancestral host hypothesis. Evidence from pheromone chemistry, dacine mating behaviour and the functional significance of lures both support and contradict the sexual selection hypothesis. Lures appear to have different proximate functions in different dacine species. Considerably greater research is needed to clarify the functional role of phytochemical lures to dacine fruit flies. The two prevalent hypotheses should be investigated independently. Specific research on dacine phylogeny and distribution of lures in plants in relation to ecological roles played by adult dacines is required to elucidate the ultimate roles of the chemicals. Exploration of female response to lures and the behavioural consequences of dacine response to these chemicals to both the insect and plant may shed light on the proximate functions of these chemicals.

Animals↗

Breeding plumage honestly signals likelihood of tapeworm infestation in females of a long-distance migrating shorebird, the bar-tailed godwit.

The indicator mechanism for sexual selection proposed by Hamilton and Zuk (i.e. that sexually selected ornaments signal parasite resistance) has received rather little observational support, and none in the case of long-distance migrant birds. Here we present a test by examining the association between helminth infestations and breeding plumage quality in bar-tailed godwits Limosa lapponica taymyrensis during their spring staging period in the Wadden Sea, The Netherlands. After a non-stop flight from West Africa, these shorebirds refuel in the Wadden Sea in preparation for a second flight to the central Siberian Arctic breeding grounds. Earlier studies have shown that only relatively heavy and well ornamented birds carry out a "top-up" moult during stopover, in which part of the contour feathers recently grown in West Africa are replaced by even fresher ones. Active body moult was therefore taken as the primary indicator of ornament quality. Of 78 birds collected between 1992 and 1997, 42% carried helminths, including four species of digenean trematodes (flukes), three species of cestodes (tapeworms) and an acanthocephalan (spiny-headed worm). Faecal samples examined for helminth eggs in another 92 birds in 1998 and 2000 showed similar rates of infestation. Actively moulting bar-tailed godwits were confirmed to be heavier and to show more extensive breeding plumage than non-moulting birds. In females, but not in males, active moult was associated with fewer cestodes and acanthocephalans. Also, breeding plumage and presence of cestodes were negatively associated in females. We argue that the quality of the breeding plumage reliably indicates parasite resistance in female godwits. The repeatability of plumage scores of females between years is consistent with such resistance having a heritable component. In contrast, male ornaments may demonstrate other qualities, e.g. an ability to combine adequate fuelling and flight performances with moult during the time-stress of migration.

Journal Article↗

Post-mating clutch piracy in an amphibian.

Female multiple mating and alternative mating systems can decrease the opportunity for sexual selection. Sperm competition is often the outcome of females mating with multiple males and has been observed in many animals, and alternative reproductive systems are widespread among species with external fertilization and parental care. Multiple paternity without associated complex behaviour related to mating or parental care is also seen in simultaneously spawning amphibians and fishes that release gametes into water. Here we report 'clutch piracy' in a montane population of the common frog Rana temporaria, a reproductive behaviour previously unknown in vertebrates with external fertilization. Males of this species clasp the females and the pair deposits one spherical clutch of eggs. No parental care is provided. 'Pirate' males search for freshly laid clutches, clasp them as they would do a female and fertilize the eggs that were left unfertilized by the 'parental' male. This behaviour does not seem to be size-dependent, and some males mate with a female and perform clutch piracy in the same season. Piracy affected 84% of the clutches and in some cases increased the proportion of eggs fertilized, providing direct fitness benefits both for the pirate males and the females. Sexual selection--probably caused by a strong male-biased sex ratio--occurs in this population, as indicated by size-assortative mating; however, clutch piracy may reduce its impact. This provides a good model to explore how alternative mating strategies can affect the intensity of sexual selection.

Animals↗

The influence of sexual and larval selection on the maintenance of polymorphism at the sepia locus in Drosophila melanogaster.

Sexual selection is measured between two strains of Drosophila melanogaster: a wild strain and a strain mutant at the sepia locus. Frequency-dependent male mating was found to successful, whereas the female genotype exerted no influence. The rarer the male genotype becomes, the greater is its mating success. A selection model is build for this behavior characteristic in which selection operates differently in the two sexes. The genetic consequencies of this odel upon the maintenance of genetic polymorphism at the sepia locus are compared to experimental data from previous population cage studies. The fit obtained with this sexual selection model is compared to that of the larval selection model previously investigated. A model composed of both sexual and larval components of fitness is presented. The role that each major selection component is expected to play in experimental populations as the gene frequency changes is discussed. Sexual selection leads to an equilibrium level higher than larval selection, and the combined model is very close to the experimental values.

Animals↗

Ornament evolution in dragon lizards: multiple gains and widespread losses reveal a complex history of evolutionary change.

The expression in females of ornaments thought to be the target of sexual selection in males is a long-standing puzzle. Two main hypotheses are proposed to account for the existence of conspicuous ornaments in both sexes (mutual ornamentation): genetic correlation between the sexes and sexual selection on females as well as males. We examined the pattern of ornament gains and losses in 240 species of dragon lizards (Agamidae) in order to elucidate the relative contribution of these two factors in the evolution of mutual ornamentation. In addition, we tested whether the type of shelter used by lizards to avoid predators predicts the evolutionary loss or constraint of ornament expression. We found evidence that the origin of female ornaments is broadly consistent with the predictions of the genetic correlation hypothesis. Ornaments appear congruently in both sexes with some lineages subsequently evolving male biased sexual dimorphism, apparently through the process of natural selection for reduced ornamentation in females. Nevertheless, ornaments have also frequently evolved in both sexes independently. This suggests that genetic correlations are potentially weak for several lineages and sexual selection on females is responsible for at least some evolutionary change in this group. Unexpectedly, we found that the evolutionary loss of some ornaments is concentrated more in males than females and this trend cannot be fully explained by our measures of natural selection.

Animals↗

Factors predicting selection of sexually violent predators for civil commitment.

This study investigated the degree to which independent variables predicted civil commitment selection in a sample of 450 sexual offenders evaluated for civil commitment as "sexually violent predators" under Florida's Jimmy Ryce Act. Using logistic regression, this study examined the relationship between the dependent variable, commitment recommendation, and several sets of independent variables. Results revealed that the statistically significant predictors of recommendations for sex offender civil commitment were diagnoses of pedophilia and paraphilia not otherwise specified (NOS), psychopathy, actuarial risk assessment scores, younger age of victim, and nonminority race (R2 = .88). Discriminant function analysis confirmed that these variables correctly predicted commitment recommendations in 90% of cases. Sex offenders recommended for commitment consistently met the criteria set forth by the U.S. Supreme Court in Kansas v. Hendricks (1997): They suffered from a mental abnormality predisposing them to sexual violence, and risk assessment determined that they were likely to reoffend.

Adult↗

Sexual isolation evolves faster than hybrid inviability in a diverse and sexually dimorphic genus of fish (Percidae: Etheostoma).

Theory predicts that sexual (or behavioral) isolation will be the first form of reproductive isolation to evolve in lineages characterized by sexual selection. Here I directly compare the rate of evolution of sexual isolation with that of hybrid inviability in a diverse and sexually dimorphic genus of freshwater fish. The magnitude of both sexual isolation and hybrid inviability were quantified for multiple pairs of allopatric species. Rates of evolution were inferred by comparing genetic distances of these species pairs with the magnitude of each form of reproductive isolation: the slope of the regression of genetic distance on the magnitude of reproductive isolation represents the rate of evolution. Of the two forms of isolation, the magnitude of sexual isolation exhibited the steeper slope of regression, indicating that sexual isolation will tend to evolve to completion earlier than hybrid inviability, strictly as a by-product of evolution in geographically isolated populations. Additional evidence from the literature is used to qualitatively compare rates of evolution of sexual isolation with that of other forms of reproductive isolation. Preliminary comparisons support the prediction that sexual isolation will evolve more rapidly than other forms. Because Etheostoma is characterized by striking sexual dimorphism, these results are consistent with the hypothesis that sexual selection for exaggerated mate-recognition characters causes the relatively rapid evolution of sexual isolation.

Animals↗

Do sexual ornaments demonstrate heightened condition-dependent expression as predicted by the handicap hypothesis?

The handicap hypothesis of sexual selection predicts that sexual ornaments have evolved heightened condition-dependent expression. The prediction has only recently been subject to experimental investigation. Many of the experiments are of limited value as they: (i) fail to compare condition dependence in sexual ornaments with suitable non-sexual trait controls; (ii) do not adequately account for body size variation; and (iii) typically consider no stress and extreme stress manipulations rather than a range of stresses similar to those experienced in nature. There is also a dearth of experimental studies investigating the genetic basis of condition dependence. Despite the common claim that sexual ornaments are condition-dependent, the unexpected conclusion from our literature review is that there is little support from well-designed experiments.

Animals↗

Developmental stress selectively affects the song control nucleus HVC in the zebra finch.

Songbirds sing complex songs as a result of evolution through sexual selection. The evolution of such sexually selected traits requires genetic control, as well as selection on their expression. Song is controlled by a discrete neural pathway in the brain, and song complexity has been shown to correlate with the volume of specific song control nuclei. As such, the development of these nuclei, in particular the high vocal centre (HVC), is thought to be the mechanism controlling signal expression indicating male quality. We tested the hypothesis that early developmental stress selectively affects adult HVC size, compared with other brain nuclei. We did this by raising cross-fostered zebra finches (Taeniopygia guttata) under stressed and controlled conditions and determining the effect on adult HVC size. Our results confirm the strong influence of environmental conditions, particularly on HVC development, and therefore on the expression of complex songs. The results also show that both environmental and genetic factors affect the development of several brain nuclei, highlighting the developmental plasticity of the songbird brain. In all, these results explain how the complex song repertoires of songbirds can evolve as honest indicators of male quality.

Analysis of Variance↗

An experimental demonstration of Fisher's principle: evolution of sexual proportion by natural selection.

Most sexually reproducing species have sexual proportions around 1:1. This major biological phenomenon remained unexplained until 1930, when FISHER proposed that it results from a mechanism of natural selection. Here we report the first experimental test of his model that obeys all its assumptions. We used a naturally occurring X-Y meiotic drive system--the sex-ratio trait of Drosophila mediopunctat--to generate female-biased experimental populations. As predicted by FISHER, these populations evolved toward equal sex proportions due to natural selection, by accumulation of autosomal alleles that direct the parental reproductive effort toward the rare sex. Classical Fisherian evolution is a rather slow mechanism: despite a very large amount of genetic variability, the experimental populations evolved from 16% of males to 32% of males in 49 generations and would take 330 generations (29 years) to reach 49%. This slowness has important implications for species potentially endangered by skewed sexual proportions, such as reptiles with temperature sex determination.

Alleles↗

Mu-, delta-, and kappa-opioid receptor agonists selectively modulate sexual behaviors in the female rat: differential dependence on progesterone.

Previous studies suggested that opioid receptor agonists infused into the lateral ventricles can inhibit (through mu receptors) or facilitate (through delta receptors) the lordosis behavior of ovariectomized (OVX) rats treated with estrogen and a low dose of progesterone. The present study investigated the behavioral and hormonal specificity of those effects using more selective opioid receptor agonists. Sexually experienced OVX rats were implanted stereotaxically with guide cannulae aimed at the right lateral ventricle. One group of rats was treated with estradiol benzoate (EB, 10 micrograms) 48 hr and progesterone (P, 250 micrograms) 4 hr before testing, whereas the other group was treated with EB alone. Rats were infused with different doses of the selective mu-receptor agonist DAMGO, the selective delta-receptor agonist DPDPE, or the selective kappa-receptor agonist U50-488. The females were placed with a sexually vigorous male in a bilevel chamber (Mendelson and Gorzalka, 1987) for three tests of sexual behavior, beginning 15, 30, and 60 min after each infusion. DAMGO reduced lordosis quotients and magnitudes significantly in rats treated with EB and P, but not in rats treated with EB alone. In contrast, DPDPE and U50-488H increased lordosis quotients and magnitudes significantly in both steroid-treatment groups. Surprisingly, measures of proceptivity, rejection responses, and level changes were not affected significantly by mu or kappa agonists, although proceptivity and rejection responses were affected by DPDPE treatment. These results suggest that the effects of lateral ventricular infusions of opioid receptor agonists on the sexual behavior of female rats are relatively specific to lordosis behavior. Moreover, the facilitation of lordosis behavior by delta- or kappa-receptor agonists is independent of progesterone treatment, whereas the inhibitory effect of mu-receptor agonists on lordosis behavior may require the presence of progesterone.

Animals↗

Male-male competition and large size mating advantage in European earwigs, Forficula auricularia.

European earwigs are sexually dimorphic in forceps shape and length. Male forceps are thought to be weapons in male contests for access to females, but recent findings suggest that females choose males on the basis of their forceps length. I investigated sexual selection on forceps length and body size and the occurrence of male-male competition. When I controlled for forceps length experimentally and statistically, relatively heavy males had greater copulation success than relatively light males. When I controlled for body size, males with relatively longer forceps had no tendency for greater copulation success than males with shorter forceps. Relatively heavy males more often took over copulations from smaller males than vice versa. Male contests were important for the outcome of mate competition, as males commonly interrupted and took over copulations. My results therefore suggest that intrasexual selection is significant in competition for copulations in male earwigs, and acts on body size. This contrasts with previous findings, which have shown intersexual selection on forceps length to be important. However, both modes of sexual selection may be acting through a two-stage process, where male-male competition first determines which males have access to females, and then through female choice among available males. Morphological measurements supported the conclusion that forceps length and body size are male secondary sexual characters, as these characters had large variance and skewed distributions in males, but were normally distributed in females. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Sexual size dimorphism in anurans.

Several hypotheses have been proposed to explain the direction and extent of sexual size dimorphism in anurans (in which males are usually smaller than females) as a result of sexual selection. Here, we present an analysis to test the hypothesis that sexual dimorphism in anurans is largely a function of differences between the sexes in life-history strategies. Morphological and demographic data for anurans were collected from the literature, and the mean size and age in each sex were calculated for 51 populations, across 30 species and eight genera. Comparisons across 14 Rana species, eight Bufo species and across the genera showed a highly significant relationship between size dimorphism, measured using the female-male size ratio, and mean female-male age difference. A comparison of a subset of 17 of these species for which phylogenetic information was available, using the method of independent contrasts, yielded a similar result. These results indicate that most of the variation in size dimorphism in the anura can be explained in terms of differences in the age structure between the sexes in breeding populations. If sexual selection has an effect on size dimorphism in anurans, it is likely to be only a secondary one.

Age Factors↗