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.     

Could sexual selection have made us psychological altruists?

Psychological altruism (being motivated by the needs of others) has a tendency to produce behaviour that is costly in evolutionary terms. How, then, could the capacity for psychological altruism evolve? One suggestion is that it is the result of sexual selection. There are, however, two problems that face such an account: first, it is not clear that the resulting behaviour would be altruistic in the relevant sense, and second, it does not seem to fit with key features of our actual helping behaviour. I will argue that both of these problems can be avoided if we adopt a modular account of desire formation.     

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.     

Is sexual selection beneficial during adaptation to environmental change?

The role of sexual selection in adaptation is disputed. A balance between sexual and viability selection can be achieved in stable environments, but environmental perturbations could change the costs and benefits arising from sexual selection and influence the rate of adaptation. Here we synthesise theoretical and empirical work on the role of sexual selection in adaptation to changed conditions. Contrasting results have been gained, but the majority of studies suggest that sexual selection has no significant effect or a negative effect on the rate of adaptation. However, once sexually selected traits start to evolve, sexual selection can accelerate adaptation. The role of sexual selection in extinction appears to be minor, but the results could be skewed.     

Are traits that experience reinforcement also under sexual selection?

Where closely related species occur in sympatry, reinforcement may result in the evolution of traits involved in species recognition that are at the same time used for within-species mate choice. Drosophila serrata lives in forested habitat on the east coast of Australia, and over the northern half of its distribution it coexists with a closely related species, Drosophila birchii. Here we show that the strength of reinforcing selection in natural populations is sufficient to generate reproductive character displacement along a 36-km transect across the contact between sympatric and allopatric populations of D. serrata. The sympatric and allopatric populations display genetically based differences in male cuticular hydrocarbons (CHCs), while female CHCs changed with latitude across the contact. The directional changes observed in male CHCs between sympatric and allopatric regions were the same changes that were generated by experimental sympatry in the laboratory, providing direct evidence that the changes across the contact zone are due to the presence of D. birchii. We show that sympatric and allopatric females differ in preference for male CHCs and that females from allopatric populations prefer allopatric-like male CHCs over sympatric-like CHCs. Male attractiveness within D. serrata may therefore be compromised by reinforcing selection, preventing the spread of sympatric-like blends to the area of allopatry.     

Intrasexual competition in females: evidence for sexual selection?

In spite of recent interest in sexual selection in females, debate exists over whether traits that influence female-female competition are sexually selected. This review uses female-female aggressive behavior as a model behavioral trait for understanding the evolutionary mechanisms promoting intrasexual competition, focusing especially on sexual selection. I employ a broad definition of sexual selection, whereby traits that influence competition for mates are sexually selected, whereas those that directly influence fecundity or offspring survival are naturally selected. Drawing examples from across animal taxa, including humans, I examine 4 predictions about female intrasexual competition based on the abundance of resources, the availability of males, and the direct or indirect benefits those males provide. These patterns reveal a key sex difference in sexual selection: Although females may compete for the number of mates, they appear to compete more so for access to high-quality mates that provide direct and indirect (genetic) benefits. As is the case in males, intrasexual selection in females also includes competition for essential resources required for access to mates. If mate quality affects the magnitude of mating success, then restricting sexual selection to competition for quantity of mates may ignore important components of fitness in females and underestimate the role of sexual selection in shaping female phenotype. In the future, understanding sex differences in sexual selection will require further exploration of the extent of mutual intrasexual competition and the incorporation of quality of mating success into the study of sexual selection in both sexes.     

Is there a role for amplifiers in sexual selection?

The amplifier hypothesis states that selection could favour the evolution of traits in signallers that improve the ability of receivers to extract honest information from other signals or cues. We provide a formal definition of amplifiers based on the receiver's mechanisms of signal perception and we present a game-theoretical model in which males advertise their quality and females use sequential-sampling tactics to choose among prospective mates. The main effect of an amplifier on the female mating strategy is to increase her mating threshold, making the female more selective as the effectiveness of the amplifier increases. The effects of the amplifier on male advertising strategy depends both on the context and on the types of the amplifier involved. We consider two different contexts for the evolution of amplifiers (when the effect of amplifiers is on signals and when it is on cues) and two types of amplifiers (the ‘neutral amplifier’, when it improves quality assessment without altering male attractiveness, and the ‘attractive amplifier’, when it improves both quality assessment and male attractiveness). The game-theoretical model provides two main results. First, neutral and attractive amplifiers represent, respectively, a conditional and an unconditional signalling strategy. In fact, at the equilibrium, neutral amplifiers are displayed only by males whose advertising level lays above the female acceptance threshold, whereas attractive amplifiers are displayed by all signalling males, independent of their quality. Second, amplifiers of signals increase the differences in advertising levels between amplifying and not-amplifying males, but they decrease the differences within each group, so that the system converges towards an ‘all-or-nothing’ signalling strategy. By applying concepts from information theory, we show that the increase in information transfer at the perception level due to the amplifier of signals is contrasted by a decrease in information transfer at the emitter level due to the increased stereotypy of male advertising strategy.     

How do natural and sexual selection contribute to sympatric speciation?

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

Sexual dimorphism in snake tail length: sexual selection,natural selection,or morphological constraint?

Male snakes typically have longer tails relative to body length than females, but the extent of this dimorphism varies among species. Three hypotheses have been suggested to explain tail dimorphism. The Morphological Constraint Hypothesis proposes that males have relatively longer tails to accommodate hemipenes and retractor muscles. The Female Reproductive Output Hypothesis proposes that females have relatively shorter tails as a secondary result of natural selection for increased reproductive capacity. The Male Mating Ability Hypothesis proposes that sexual selection favours relatively longer tails in males during courtship. These hypotheses make different predictions about the relationships among tail length, body size, male reproductive morphology, female reproductive output, mode of reproduction, and male mating behaviour among and within taxa. Predictions were tested using published data for 56 genera in the family Colubridae and original data for the water snake, Nerodia sipedon. Tail length dimorphism was more male-biased in tam having relatively short tails (r=–0.52, P < 0.001), hemipenes and retractor muscles occupied a greater proportion of the tail in taxa having relatively short tails (r=– 0.71, P < 0.00l and r=– 0.66, P = 0.001, respectively), and tail length dimorphism was more male-biased in taxa in which body size dimorphism was more female-biased (r=– 0.60, P < 0.001). These results support both the Morphological Constraint Hypotheses and the Female Reproductive Output Hypothesis. However, tests of other predictions, including those regarding patterns within N. sipedon , failed to support any of the three hypotheses. Comparisons among taxa suggest several species in which further tests of these hypotheses would be especially appropriate.     

Simple signaling games of sexual selection (Grafen’s revisited)

We investigate several versions of a simple game of sexual selection, to explore the role of secondary sexual characters (the “handicap paradox”) with the tools of signaling theory. Our models admit closed form solutions. They are very much inspired by Grafen’s (J Theor Biol 144:517–546, 1990a; J Theor Biol 144:473–516, 1990b) seminal companion papers. By merging and simplifying his two approaches, we identify a not so minor artifact in the seminal study. We propose an alternative model to start with Grafen’s sexual selection theory, with several similarities with Getty (Anim Behav 56:127–130, 1998).     

Trilobite spines and beetle horns: sexual selection in the Palaeozoic?

Raphiophorid trilobites commonly bore median cephalic protuberances such as spines or bulbs, showing a remarkable variety of form. It is unlikely that their primary function was for protection or in hydrodynamics. A case is made that they were secondary sexual features, by comparison with similar morphological structures developed on rhinoceros beetles and other arthropods. This interpretation is supported by four lines of evidence: their ontogeny, their diversity, the existence of plausible examples of sexual dimorphs in some cases and the fact that they show positive allometry.     

The retrolateral tibial apophysis in spiders—shaped by sexual selection?

The functional significance of the retrolateral tibial apophysis (rta) on the male pedipalps in four spider species with different mating positions is investigated with the help of histological serial sections prepared after freeze-fixing copulating pairs with liquid nitrogen. The results of this study, as well as most data in previous works, suggest that the rta is mostly used to fix the male pedipalp to the female epigyne in order to ensure the intromission of the sperm transferring embolus. This is in accordance with the female choice hypothesis on genitalia which predicts that species-specific genital structures should directly or indirectly contact the female during copulation and thus be shaped by sexual selection.     

Sexually selected sexual selection: Can evolutionary retribution explain female ornamental colour?

By preferring mates with increasingly costly ornaments or courtship displays, females cause an escalation of male reproductive costs. Such increased costs should promote male selectivity based on fecundity‐linked female attributes, leading to female ornamentation in species with traditional sex roles. Consequently, female ornamentation should evolve more frequently in taxa where male reproduction is costly than in comparable taxa where it is cheaper. We assessed the prevalence of female ornamental colouration in two clades of viviparous cyprinodontid fish: the Goodeinae, where stringent female choice imposes male mating costs, and the Poeciliinae, whose males can circumvent female mate choice. We found that although in the Poeciliinae female ornamental colour is a correlated, but paler version of male coloration, females of the Goodeinae often display vivid ornamental colours that are distinct from those of males. Thus, male and female ornaments are not (phylo)genetically correlated in the Goodeinae. Furthermore, phylogenetic signal on male and female colour is clearly detectable in the Poeciliinae, but absent in the Goodeinae, suggesting that ornamental colour of males and females in the latter may be the consequence of selection. Given that enforceable female choice has promoted male ornaments, we propose that evolutionary retribution has promoted distinct female ornaments in the Goodeinae.     

Can social and sexual selection explain the bizarre snout of proterosuchid archosauriforms?

Proterosuchids are a clade of quadrupedal, carnivorous Permo-Triassic diapsids crucial to understand the successful evolutionary radiation of archosaurs during the Mesozoic. The importance and good fossil record of proterosuchids nourished a renewed interest in recent years, but no function has been proposed for their bizarre snouts. An oversized and downturned premaxilla with up to nine teeth with continuous replacement is present in all proterosuchid species and seems to have represented a physiologically costly phenotype that increased towards adulthood. A non-functional or a species recognition hypothesis are not supported as evolutionary mechanisms that drove this phenotype because features expected for these explanations tend to have a very low or zero physiological cost. There is no evidence favouring – but neither rejecting ? that this morphology can be explained by non-sexual and non-social natural selection alone. Mutual social and/or sexual selection is favoured here as the most unambiguously supported explanation for the function and origin of the bizarre snout of proterosuchids based on several lines of evidence, including costliness, positive allometry, positive changes in growth rates and modern analogues. Social and/or sexual selection may have been important evolutionary mechanism in the dawn of the lineage that gave rise to crocodiles and dinosaurs.     

Alternative phenotypes and sexual selection: can dichotomous handicaps honestly signal quality?

Considerable theoretical and empirical effort has been focused on the potential of continuously variable sexual traits to honestly indicate male quality, but relatively little effort has been devoted to a similar evolutionary role for dimorphic traits. Male dimorphisms, associated with conditionally expressed alternative reproductive tactics, represent extreme phenotypic plasticity. Evidence suggests that considerable heritable variation exists in the 'liability' underlying many threshold traits; if this liability is correlated with the genetic quality of males, dimorphic traits have the potential to be reliable indicators. We investigated the genetic architecture of phenotypically plastic morph expression in the context of condition-dependent signalling theory. Male morph in the mite Sancassania berlesei is condition dependent: 'fighters' armed with thickened and sharp third pairs of legs emerge from heavier nymphs than unarmoured 'scramblers'. We selected on male morph in three replicate 'fighter' and 'scrambler' lines and recorded a significant response to selection over seven generations; this was due to a shift in the threshold reaction norm but the lines showed no correlated response in condition. This is inconsistent with models predicting a substantial genetic correlation between condition and sexual trait expression. We discuss why dimorphic sexual traits may show more condition-independent genetic variance than continuous sexual traits.     

Is sexual selection driving diversification of the bioluminescent ponyfishes (Teleostei: Leiognathidae)?

Sexual selection may facilitate genetic isolation among populations and result in increased rates of diversification. As a mechanism driving diversification, sexual selection has been invoked and upheld in numerous empirical studies across disparate taxa, including birds, plants and spiders. In this study, we investigate the potential impact of sexual selection on the tempo and mode of ponyfish evolution. Ponyfishes (Leiognathidae) are bioluminescent marine fishes that exhibit sexually dimorphic features of their unique light-organ system (LOS). Although sexual selection is widely considered to be the driving force behind ponyfish speciation, this hypothesis has never been formally tested. Given that some leiognathid species have a sexually dimorphic LOS, whereas others do not, this family provides an excellent system within which to study the potential role of sexual selection in diversification and morphological differentiation. In this study, we estimate the phylogenetic relationships and divergence times for Leiognathidae, investigate the tempo and mode of ponyfish diversification, and explore morphological shape disparity among leiognathid clades. We recover strong support for a monophyletic Leiognathidae and estimate that all major ponyfish lineages evolved during the Paleogene. Our studies of ponyfish diversification demonstrate that there is no conclusive evidence that sexually dimorphic clades are significantly more species rich than nonsexually dimorphic lineages and that evidence is lacking to support any significant diversification rate increases within ponyfishes. Further, we detected a lineage-through-time signal indicating that ponyfishes have continuously diversified through time, which is in contrast to many recent diversification studies that identify lineage-through-time patterns that support mechanisms of density-dependent speciation. Additionally, there is no evidence of sexual selection hindering morphological diversity, as sexually dimorphic taxa are shown to be more disparate in overall shape morphology than nonsexually dimorphic taxa. Our results suggest that if sexual selection is occurring in ponyfish evolution, it is likely acting only as a genetic isolating mechanism that has allowed ponyfishes to continuously diversify over time, with no overall impact on increases in diversification rate or morphological disparity.     

Can sexual selection drive female life histories? A comparative study on Galliform birds

Sexual selection has been identified as a major evolutionary force shaping male life history traits but its impact on female life history evolution is less clear. Here we examine the impact of sexual selection on three key female traits (body size, egg size and clutch size) in Galliform birds. Using comparative independent contrast analyses and directional discrete analyses, based on published data and a new genera-level supertree phylogeny of Galliform birds, we investigated how sexual selection [quantified as sexual size dimorphism (SSD) and social mating system (MS)] affects these three important female traits. We found that female body mass was strongly and positively correlated with egg size but not with clutch size, and that clutch size decreased as egg size increased. We established that SSD was related to MS, and then used SSD as a proxy of the strength of sexual selection. We found both a positive relationship between SSD and female body mass and egg size and that increases in female body mass and egg size tend to occur following increases in SSD in this bird order. This pattern of female body mass increases lagging behind changes in SSD, established using our directional discrete analysis, suggests that female body mass increases as a response to increases in the level of sexual selection and not simply through a strong genetic relationship with male body mass. This suggests that sexual selection is linked to changes in female life history traits in Galliformes and we discuss how this link may shape patterns of life history variation among species.