The evolution of sexual dimorphism in parasitic cuckoos: sexual selection or coevolution?

Sexual dimorphism is ubiquitous in animals and can result from selection pressure on one or both sexes. Sexual selection has become the predominant explanation for the evolution of sexual dimorphism, with strong selection on size-related mating success in males being the most common situation. The cuckoos (family Cuculidae) provide an exceptional case in which both sexes of many species are freed from the burden of parental care but where coevolution between parasitic cuckoos and their hosts also results in intense selection. Here, we show that size and plumage differences between the sexes in parasitic cuckoos are more likely the result of coevolution than sexual selection. While both sexes changed in size as brood parasitism evolved, we find no evidence for selection on males to become larger. Rather, our analysis indicates stronger selection on parasitic females to become smaller, resulting in a shift from dimorphism with larger females in cuckoos with parental care to dimorphism with larger males in parasitic species. In addition, the evolution of brood parasitism was associated with more cryptic plumage in both sexes, but especially in females, a result that contrasts with the strong plumage dimorphism seen in some other parasitic birds. Examination of the three independent origins of brood parasitism suggests that different parasitic cuckoo lineages followed divergent evolutionary pathways to successful brood parasitism. These results argue for the powerful role of parasite-host coevolution in shaping cuckoo life histories in general and sexual dimorphism in particular.     

Testosterone in females: mediator of adaptive traits, constraint on sexual dimorphism, or both?

When selection on males and females differs, the sexes may diverge in phenotype. Hormones serve as a proximate regulator of sex differences by mediating sex-biased trait expression. To integrate these perspectives, we consider how suites of traits mediated by the same hormone in both sexes might respond to selection. In male birds, plasma testosterone (T) varies seasonally and among species according to mating system. When elevated experimentally, it is known to enhance some components of fitness and to decrease others. We report that female T also varies seasonally and co-varies with male T. Female T is higher in relation to male T in sexually monomorphic species and is higher absolutely in females of species with socially monogamous mating systems, which suggests adaptation. We also consider the effect of experimentally elevated T on females and whether traits are sensitive to altered T. We hypothesize that sensitive traits could become subject to selection after a natural change in T and that traits with opposing fitness consequences in males and females could constrain dimorphism. Results from birds, including the dark-eyed junco (Junco hyemalis), reveal many sensitive traits, some of which appear costly and may help to account for observed levels of sexual dimorphism.     

Sexual dimorphism and interpopulation differences in lizard hind limb length: locomotor performance or chemical signalling?

Intraspecific variation in morphology has often been related to fitness differences through its effects on performance. In lizards, variation in hind limb length can be shaped by natural selection for increased locomotor performance, sexual selection on the number or size of femoral pores involved in chemical signalling, or both. Here, we analyse the selective forces involved in sexual dimorphism and differences in hind limb length between two populations of Psammodromus algirus living at different elevation. Males were more robust and had longer hind limbs and limb segments than females, and low‐elevation lizards had longer limbs than high‐elevation lizards. However, differences in locomotor performance were small and non‐significant, making natural selection for faster runs an unlikely explanation for the observed pattern. On the other hand, males had more femoral pores than females, and lizards had more pores at lower elevation, although the difference was significant only for males (which invest more in chemical signalling). In males, the number of pores, which remains constant along a lizard's life, was not correlated with hind limb length. However, femur length was positively correlated with mean pore size, allowing low‐elevation males to have larger than expected pores, which could increase the effectiveness with which they spread their signals in a dry and warm habitat where chemicals become volatile rapidly. Also, saturation of the sexual coloration of the head was higher for low‐elevation males, suggesting that sexual selection pressures may be more intense. Overall, our results indicate that sexual selection plays a significant role in shaping intraspecific variation in hind limb length. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 318–329.     

Dimorphic male midshipman fish: reduced sexual selection or sexual selection for reduced characters?

In most taxa with male dimorphisms, some males are large inbody size with exaggerated secondary sexual characters (exaggeratedmorph), whereas other males in the same population are smalland have reduced secondary sexual characters (reduced morph).What selective pressures cause male dimorphisms? Reduced morphologiesmay result when a) some males develop a morphology that, inthe absence of sexual selection pressures for an exaggeratedmorphology, reduces energetic and developmental costs and/orb) some males opt for an alternative morphology that does wellat an alternative behavioral tactic such as cuckoldry. The 2mechanisms could act together, but each alone is theoreticallysufficient to drive dimorphisms. Here, we tested hypothesis\"b\" (sexual selection for reduced characters) in the plainfinmidshipman fish, Porichthys notatus. Behavioral plasticity betweenterritoriality and cuckoldry in an exaggerated male morph (typeI) allows for a direct comparison of cuckoldry by exaggeratedmorph males to cuckoldry by reduced morph (type II) males. Comparedwith type I cuckolders, type II cuckolders were able to remainnear the nest for longer periods before being chased by theterritorial type I male, suggesting that the reduced type IImorphology allows type II males to prolong the time before attackby territorial males. Combined with other studies showing arole of sexual selection in maintaining the exaggerated morph,the data support the \"sexual selection for reduced characters\"hypothesis and elucidate how sexual selection can act in differentways on different males to maintain 2 male morphologies withina single species.     

Species richness in agamid lizards: chance,body size,sexual selection or ecology?

Why does species richness vary so greatly across lineages? Traditionally, variation in species richness has been attributed to deterministic processes, although it is equally plausible that it may result from purely stochastic processes. We show that, based on the best available phylogenetic hypothesis, the pattern of cladogenesis among agamid lizards is not consistent with a random model, with some lineages having more species, and others fewer species, than expected by chance. We then use phylogenetic comparative methods to test six types of deterministic explanation for variation in species richness: body size, life history, sexual selection, ecological generalism, range size and latitude. Of eight variables we tested, only sexual size dimorphism and sexual dichromatism predicted species richness. Increases in species richness are associated with increases in sexual dichromatism but reductions in sexual size dimorphism. Consistent with recent comparative studies, we find no evidence that species richness is associated with small body size or high fecundity. Equally, we find no evidence that species richness covaries with ecological generalism, latitude or range size.     

Is natural selection promoting sexual dimorphism in the Green-backed Firecrown Hummingbird ( Sephanoides Sephaniodes )?

In many hummingbird species there is an opposite pattern of sexual dimorphism in bill length and other morphometric measures of body size. These differences seem to be closely related with differences in foraging ecology directly associated with a different resource exploitation strategy. The aim of this study was to assess if natural selection is acting on wing length and bill size in hummingbird males and females with different resource exploitation strategies (i.e., territorial males and non-territorial females). If competition for resources promotes sexual dimorphism as a selective pressure, males should be subjected to negative directional selection pressure for wing length and no selection pressure over bill size, while females should undergo positive directional selection pressure for both bill size and wing length. The morphometric data we collected suggests that there is no selection for wing length and bill size in male hummingbirds. In contrast, our females exhibited positive directional selection for both wing length and bill size. Although we cannot reject sexual selection acting on sexually dimorphic traits, this study suggests that natural selection may promote sexual dimorphism in traits that are closely related with hummingbird foraging ecology and resource exploitation strategies.     

Colour diversification in Malawi cichlids: evidence for adaptation,reinforcement or sexual selection?

The mechanisms responsible for the explosive radiations of haplochromine cichlid fishes in Africa's great lakes remain controversial. Since species thought to be closely related often differ most apparendy in male breeding colours, I examined patterns of male colour variation in rock-dwelling cichlids from Lake Malawi to test whether initial divergence between species is likely to have been caused by adaptation to differing habitats, by selection against hybridization (reinforcement or character displacement), or by sexual selection. I found that all significant variation in overall male colour occurs within, not between, species complexes, in contrast to variation in habitat and behavioural traits, which differed significandy between complexes. Male colour does not vary significandy with habitat characteristics such as water depth, and sympatric species differ no more in colour than allopatric ones. These results fail to provide support for adaptation or reinforcement as initial causes of colour divergence, and are consistent with the possibility that divergence may instead have been caused by sexual selection, but a direct test of sexual selection is not yet possible.     

Sexual dimorphism in a Lower Miocene moronid?

A series of questionable elements in certain specimens ofMorone cf.aequalis (Koken 1891) from Lower Miocene deposits near the village of Berkersheim, N of Frankfurt a. M. (Hessen, Germany) is described, which has not been known from any other percoid before. These elements are fully ossified and cover the cheek and the preopercular region. Even within well-preserved material, they are only present in some specimens. Therefore, they may be specialized structures that are indicative for sexual dimorphism. Nevertheless, they clearly differ from all other respective structures that have been described from teleosts: Multicellular epidermal horny tubercles (“breeding tubercles”) mainly consist of keratine and not of calciumphosphate. By contrast, contact organs consist of bone and are located mainly at the surface of the fin rays and scales, respectively. At present, “breeding tubercles” are the favorite interpretation and the original substance may have been replaced via post-mortem phosphatization.     

Why do males emerge before females? Sexual size dimorphism drives sexual bimaturism in insects

Conspecific females and males often follow different development trajectories which leads to sex differences in age at maturity (sexual bimaturism, SBM). Whether SBM is typically selected for per se (direct selection hypothesis) or merely represents a side-effect of other sex-related adaptations (indirect selection hypothesis) is, however, still an open question. Substantial interspecific variation in the direction and degree of SBM, both in invertebrates and vertebrates, calls for multi-species studies to understand the relative importance of its evolutionary drivers. Here we use two complementary approaches to evaluate the evolutionary basis of SBM in insects. For this purpose, we assembled an extensive literature-derived data set of sex-specific development times and body sizes for a taxonomically and ecologically wide range of species. We use these data in a meta-analytic framework to evaluate support for the direct and indirect selection hypotheses. Our results confirm that protandry – males emerging as adults before females – is the prevailing form of SBM in insects. Nevertheless, protandry is not as ubiquitous as often presumed: females emerged before males (= protogyny) in about 36% of the 192 species for which we had data. Moreover, in a considerable proportion of species, the sex difference in the timing of adult emergence was negligible. In search for the evolutionary basis of SBM, we found stronger support for the hypothesis that explains SBM by indirect selection. First, across species, the direction and degree of SBM appeared to be positively associated with the direction and degree of sexual size dimorphism (SSD). This is consistent with the view that SBM is a correlative by-product of evolution towards sexually dimorphic body sizes. Second, within protandrous species, the degree of protandry typically increased with plastic increase in development time, with females prolonging their development more than males in unfavourable conditions. This pattern is in conflict with the direct selection hypothesis, which predicts the degree of protandry to be insensitive to the quality of the juvenile environment. These converging lines of evidence support the idea that, in insects, SBM is generally a by-product of SSD rather than a result of selection on the two sexes to mature at different times. It appears plausible that selective pressures on maturation time per se generally cannot compete with viability- and fecundity-mediated selection on insect body sizes. Nevertheless, exceptions certainly exist: there are undeniable cases of SBM where this trait has evolved in response to direct selection. In such cases, either the advantage of sex difference in maturation time must have been particularly large, or fitness effects of body size have been unusually weak.     

Phenotypic selection in natural populations: what limits directional selection?

Studies of phenotypic selection document directional selection in many natural populations. What factors reduce total directional selection and the cumulative evolutionary responses to selection? We combine two data sets for phenotypic selection, representing more than 4,600 distinct estimates of selection from 143 studies, to evaluate the potential roles of fitness trade-offs, indirect (correlated) selection, temporally varying selection, and stabilizing selection for reducing net directional selection and cumulative responses to selection. We detected little evidence that trade-offs among different fitness components reduced total directional selection in most study systems. Comparisons of selection gradients and selection differentials suggest that correlated selection frequently reduced total selection on size but not on other types of traits. The direction of selection on a trait often changes over time in many temporally replicated studies, but these fluctuations have limited impact in reducing cumulative directional selection in most study systems. Analyses of quadratic selection gradients indicated stabilizing selection on body size in at least some studies but provided little evidence that stabilizing selection is more common than disruptive selection for most traits or study systems. Our analyses provide little evidence that fitness trade-offs, correlated selection, or stabilizing selection strongly constrains the directional selection reported for most quantitative traits.     

Sexual differences in habitat use by small mammals: evolutionary strategy or reproductive constraint?

Summary This study examines the hypothesis that intersexual differences in habitat use by small mammals is an adaptive strategy. Specifically, I evaluate the occurrence of sexual differences in habitat by Peromyscus and Microtus, and test possible adaptive mechanisms which may lead to such differences.White-footed mice and meadow voles were live-trapped and microhabitat quantified in four habitats. Sexual differences in microhabitat use were found in three of 12 comparisons. These differences may be related to density dependent resource subdivision and habitat heterogencity, but neither of these can account completely for the observed patterns. Sexual differences in habitat do not appear to be an evolutionary strategy maximizing reproductive effort by females, or reducing predation. These sexual differences may reduce intraspecific resource overlap, or may simply reflect reproductive constraints limiting female habitat use to suitable nesting areas.     

Sexual dimorphism in birds: why are there so many different forms of dimorphism?

Variation in the extent of sexual dimorphism among bird species is traditionally attributed to differences in social mating system. However, there are many different forms of dimorphism among birds, and not all of them show an obvious correlation with social mating system. For example, recent work has shown that many highly polygamous species are, in fact, monomorphic, whereas many putatively monogamous species are dimorphic. In this paper we break up sexual dimorphism into subcomponents and then use comparative analyses to examine the pattern of covariation between these subcomponents and various aspects of sexual, social, and parental behaviour. Our first finding is that size dimorphism and plumage-colour dimorphism do not show the same pattern of covariation. Differences in size dimorphism are associated with variation in social mating system and sex differences in parental care, whereas differences in plumage-colour dimorphism are associated with variation in the frequency of extra-bond paternity. These results suggest that size dimorphism is associated with the sort of intrasexual competition described by traditional classifications of social mating system, whereas plumage-colour dimorphism is associated with cryptic female choice. However, when we break up plumage-colour dimorphism according to whether it is due to melanins, carotenoids or structural colours, we find that each category of plumage-colour dimorphism shows a different pattern of covariation. The correlation between overall plumage-colour dimorphism and the rate of extra-bond paternity is due to structural colours, whereas melanin-based dimorphism is associated with sex differences in parental care. The former result is particularly interesting given that new work suggests structural colours are associated with active sexual displays and the reflection of ultraviolet light.     

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.     

Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Trait selection in flowering plants: how does sexual selection contribute?

By highlighting and merging the frameworks of sexual selectionenvisioned by Arnold (1994) and Murphy (1998), we discuss howsexual selection can occur in plants even though individualsdo not directly interact. We review studies on traits that influencepollen export and receipt in a variety of hermaphroditic andgynodioecious plants with the underlying premise that pollinationdynamics influences mate acquisition. Most of the studies reviewedfound that phenotypes that enhance pollen export are in harmonywith those that enhance pollen receipt suggesting that in manycases pollinator visitation rates limit both male and femalefunction. In contrast, fewer traits were under opposing selection;but when they were, the traits most often were associated withenhancing the specific aspects of a given sex function. Ourreview helps clarify and illustrate why sexual selection canbe a component of trait evolution in hermaphrodite plants.     

Sexual dimorphism in immunocompetence: what does life-history theory predict?

Sexual dimorphism in immunocompetence, usually in the directionof inferior male immunocompetence, has historically been explainedas the result of proximate physiological mechanisms such asthe immunosuppressive effects of the male hormone testosterone.More recently, it has been argued that this pattern is bestunderstood as a result of resource-based trade-offs betweenmale mating effort and immune defense, a trade-off that femalesdo not make. The central prediction of this hypothesis is thatas the strength of sexual selection on males increases, themagnitude of the sex differences in immunocompetence will increase.Two implicit assumptions of this argument are that 1) longevityis of more importance for female than for male fitness and 2)that the primary benefit of immunocompetence is increased longevity.However, both of these assumptions may not be as broadly applicableas has been argued. We have modeled the optimal allocation toimmunocompetence for males and females without making theseassumptions. We find that the optimal allocation to immune defensefor males decreases as the strength of sexual selection increases,as predicted. However, males may still invest more, relativeto females, into immunocompetence if the impact of parasiteson condition differs for the sexes and/or if the relationshipbetween condition and reproduction differs for the sexes. Weargue that these previously overlooked assumptions may be criticalfor predicting sex-specific patterns of immunocompetence.     

Paracorethrura Melichar, 1915 (Hemiptera: Lophopidae): two distinct species or sexual dimorphism in a species?

A taxonomic update is provided for Paracorethrura Melichar, 1915. P. arrhen n. sp. is described from Vietnam and brings up the number of species to two for the genus: P. iocnemis (Jacobi, 1905) being known only by female specimens and P. arrhen only by males, CO1 sequences are provided in order to check if they are truly two different species or if we are witnessing some sexual dimorphism.     

Establishing sexual dimorphism: conservation amidst diversity?

The molecular mechanisms that control sexual dimorphism are very different in distantly related animals. Did sex determination arise several times with different regulatory mechanisms, or is it an ancient process with little surviving evidence of ancestral genes? The recent identification of related sexual regulators in different phyla indicates that some aspects of sexual regulation might be ancient. Studies of sex-determining mechanisms are beginning to reveal how sexual dimorphism arises and evolves.