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1.  Quantifying Variability of Avian Colours: Are Signalling Traits More Variable? 
PLoS ONE  2008;3(2):e1689.
Background
Increased variability in sexually selected ornaments, a key assumption of evolutionary theory, is thought to be maintained through condition-dependence. Condition-dependent handicap models of sexual selection predict that (a) sexually selected traits show amplified variability compared to equivalent non-sexually selected traits, and since males are usually the sexually selected sex, that (b) males are more variable than females, and (c) sexually dimorphic traits more variable than monomorphic ones. So far these predictions have only been tested for metric traits. Surprisingly, they have not been examined for bright coloration, one of the most prominent sexual traits. This omission stems from computational difficulties: different types of colours are quantified on different scales precluding the use of coefficients of variation.
Methodology/Principal Findings
Based on physiological models of avian colour vision we develop an index to quantify the degree of discriminable colour variation as it can be perceived by conspecifics. A comparison of variability in ornamental and non-ornamental colours in six bird species confirmed (a) that those coloured patches that are sexually selected or act as indicators of quality show increased chromatic variability. However, we found no support for (b) that males generally show higher levels of variability than females, or (c) that sexual dichromatism per se is associated with increased variability.
Conclusions/Significance
We show that it is currently possible to realistically estimate variability of animal colours as perceived by them, something difficult to achieve with other traits. Increased variability of known sexually-selected/quality-indicating colours in the studied species, provides support to the predictions borne from sexual selection theory but the lack of increased overall variability in males or dimorphic colours in general indicates that sexual differences might not always be shaped by similar selective forces.
doi:10.1371/journal.pone.0001689
PMCID: PMC2253496  PMID: 18301766
2.  Male genotype influences female reproductive investment in Drosophila melanogaster 
In many species, males can influence the amount of resources their mates invest in reproduction. Two favoured hypotheses for this observation are that females assess male quality during courtship or copulation and alter their investment in offspring accordingly, or that males manipulate females to invest heavily in offspring produced soon after mating. Here, we examined whether there is genetic variation for males to influence female short-term reproductive investment in Drosophila melanogaster, a species with strong sexual selection and substantial sexual conflict. We measured the fecundity and egg size of females mated to males from multiple isofemale lines collected from populations around the globe. Although these traits were not strongly influenced by the male's population of origin, we found that 22 per cent of the variation in female short-term reproductive investment was attributable to the genotype of her mate. This is the first direct evidence that male D. melanogaster vary genetically in their proximate influence on female fecundity, egg size and overall reproductive investment.
doi:10.1098/rspb.2010.2272
PMCID: PMC3107629  PMID: 21159677
maternal investment; fecundity; egg size; differential allocation hypothesis; sexual conflict; Drosophila melanogaster
3.  Condition-dependent expression of pre- and postcopulatory sexual traits in guppies 
Ecology and Evolution  2013;3(7):2197-2213.
Female choice can impose persistent directional selection on male sexually selected traits, yet such traits often exhibit high levels of phenotypic variation. One explanation for this paradox is that if sexually selected traits are costly, only the fittest males are able to acquire and allocate the resources required for their expression. Furthermore, because male condition is dependent on resource allocation, condition dependence in sexual traits is expected to underlie trade-offs between reproduction and other life-history functions. In this study we test these ideas by experimentally manipulating diet quality (carotenoid levels) and quantity in the guppy (Poecilia reticulata), a livebearing freshwater fish that is an important model for understanding relationships between pre- and post-copulatory sexually selected traits. Specifically, we test for condition dependence in the expression of pre- and postcopulatory sexual traits (behavior, ornamentation, sperm traits) and determine whether diet manipulation mediates relationships among these traits. Consistent with prior work we found a significant effect of diet quantity on the expression of both pre- and postcopulatory male traits; diet-restricted males performed fewer sexual behaviors and exhibited significant reductions in color ornamentation, sperm quality, sperm number, and sperm length than those fed ad libitum. However, contrary to our expectations, we found no significant effect of carotenoid manipulation on the expression of any of these traits, and no evidence for a trade-off in resource allocation between pre- and postcopulatory episodes of sexual selection. Our results further underscore the sensitivity of behavioral, ornamental, and ejaculate traits to dietary stress, and highlight the important role of condition dependence in maintaining the high variability in male sexual traits.
doi:10.1002/ece3.632
PMCID: PMC3728957  PMID: 23919162
Carotenoids; dietary manipulation; genetic variation; lek paradox; trade-offs
4.  Estimation of heritability, evolvability and genetic correlations of two pollen and pistil traits involved in a sexual conflict over timing of stigma receptivity in Collinsia heterophylla (Plantaginaceae) 
Annals of Botany  2012;110(1):91-99.
Background and Aims
Heritable genetic variation is crucial for selection to operate, yet there is a paucity of studies quantifying such variation in interactive male/female sexual traits, especially those of plants. Previous work on the annual plant Collinsia heterophylla, a mixed-mating species, suggests that delayed stigma receptivity is involved in a sexual conflict: pollen from certain donors fertilize ovules earlier than others at the expense of reduced maternal seed set and lower levels of pollen competition.
Methods
Parent–offspring regressions and sib analyses were performed to test for heritable genetic variation and co-variation in male and female interactive traits related to the sexual conflict.
Key Results
Some heritable variation and evolvability were found for the female trait (delayed stigma receptivity in presence of pollen), but no evidence was found for genetic variation in the male trait (ability to fertilize ovules early). The results further indicated a marginally significant correlation between a male's ability to fertilize early and early stigma receptivity in offspring. However, despite potential indirect selection of these traits, antagonistic co-evolution may not occur given the lack of heritability of the male trait.
Conclusions
To our knowledge, this is the first study of a plant or any hermaphrodite that examines patterns of genetic correlation between two interactive sexual traits, and also the first to assess heritabilities of plant traits putatively involved in a sexual conflict. It is concluded that the ability to delay fertilization in presence of pollen can respond to selection, while the pollen trait has lower evolutionary potential.
doi:10.1093/aob/mcs084
PMCID: PMC3380587  PMID: 22645118
Antagonistic trait; co-evolution; Collinsia heterophylla; evolvability; genetic correlation; genetic variation; heritability; mate choice; pollen competition; sexual conflict; stigma receptivity
5.  Family feuds: social competition and sexual conflict in complex societies 
Darwin was initially puzzled by the processes that led to ornamentation in males—what he termed sexual selection—and those that led to extreme cooperation and altruism in complex animal societies—what was later termed kin selection. Here, I explore the relationships between sexual and kin selection theory by examining how social competition for reproductive opportunities—particularly in females—and sexual conflict over mating partners are inherent and critical parts of complex altruistic societies. I argue that (i) patterns of reproductive sharing within complex societies can drive levels of social competition and reproductive conflict not only in males but also in females living in social groups, and ultimately the evolution of female traits such as ornaments and armaments; (ii) mating conflict over female choice of sexual partners can influence kin structure within groups and drive the evolution of complex societies; and (iii) patterns of reproductive sharing and conflict among females may also drive the evolution of complex societies by influencing kin structure within groups. Ultimately, complex societies exhibiting altruistic behaviour appear to have only arisen in taxa where social competition over reproductive opportunities and sexual conflict over mating partners were low. Once such societies evolved, there were important selective feedbacks on traits used to regulate and mediate intra-sexual competition over reproductive opportunities, particularly in females.
doi:10.1098/rstb.2011.0283
PMCID: PMC3391424  PMID: 22777018
kin selection; sexual selection; social selection; sexual conflict; reproductive conflict; mating conflict
6.  Postcopulatory Sexual Selection Is Associated with Reduced Variation in Sperm Morphology 
PLoS ONE  2007;2(5):e413.
Background
The evolutionary role of postcopulatory sexual selection in shaping male reproductive traits, including sperm morphology, is well documented in several taxa. However, previous studies have focused almost exclusively on the influence of sperm competition on variation among species. In this study we tested the hypothesis that intraspecific variation in sperm morphology is driven by the level of postcopulatory sexual selection in passerine birds.
Methodology/Findings
Using two proxy measures of sperm competition level, (i) relative testes size and (ii) extrapair paternity level, we found strong evidence that intermale variation in sperm morphology is negatively associated with the degree of postcopulatory sexual selection, independently of phylogeny.
Conclusions/Significance
Our results show that the role of postcopulatory sexual selection in the evolution of sperm morphology extends to an intraspecific level, reducing the variation towards what might be a species-specific ‘optimum’ sperm phenotype. This finding suggests that while postcopulatory selection is generally directional (e.g., favouring longer sperm) across avian species, it also acts as a stabilising evolutionary force within species under intense selection, resulting in reduced variation in sperm morphology traits. We discuss some potential evolutionary mechanisms for this pattern.
doi:10.1371/journal.pone.0000413
PMCID: PMC1855076  PMID: 17476335
7.  A potential resolution to the lek paradox through indirect genetic effects 
Females often prefer males with elaborate traits, even when they receive no direct benefits from their choice. In such situations, mate discrimination presumably has genetic advantages; selective females will produce offspring of higher genetic quality. Over time, persistent female preferences for elaborate secondary-sexual traits in males should erode genetic variance in these traits, eventually eliminating any benefit to the preferences. Yet, strong female preferences persist in many taxa. This puzzle is called the lek paradox and raises two primary questions: do females obtain genetic benefits for offspring by selecting males with elaborate secondary-sexual characteristics and, if so, how is the genetic variation in these male traits maintained? We suggest that indirect genetic effects may help to resolve the lek paradox. Maternal phenotypes, such as habitat selection behaviours and offspring provisioning, often influence the condition and the expression of secondary-sexual traits in sons. These maternal influences are commonly genetic based (i.e. they are indirect genetic effects). Females choosing mates with elaborate traits may receive ‘good genes’ for daughters in the form of effective maternal characteristics. Recognizing the significance of indirect genetic effects may be important to our understanding of the process and consequences of sexual selection.
doi:10.1098/rspb.2006.0413
PMCID: PMC2176171  PMID: 17341455
lek paradox; indirect genetic effects; sexual selection; condition dependence; good genes; maternal effects
8.  Does divergence in female mate choice affect male size distributions in two cave fish populations? 
Biology Letters  2008;4(5):452-454.
Sexual selection by female choice can maintain male traits that are counter selected by natural selection. Alteration of the potential for sexual selection can thus lead to shifts in the expression of male traits. We investigated female mate choice for large male body size in a fish (Poecilia mexicana) that, besides surface streams, also inhabits two caves. All four populations investigated, exhibited an ancestral visual preference for large males. However, only one of the cave populations also expressed this female preference in darkness. Hence, the lack of expression of female preference in darkness in the other cave population leads to relaxation of sexual selection for large male body size. While P. mexicana populations with size-specific female mate choice are characterized by a pronounced male size variation, the absence of female choice in one cave coincides with the absence of large bodied males in that population. Our results suggest that population differences in the potential for sexual selection may affect male trait variation.
doi:10.1098/rsbl.2008.0259
PMCID: PMC2610082  PMID: 18559308
non-visual mate choice; sensory shift; sexual selection; size variation; Poecilia mexicana (Poeciliidae)
9.  Environmental variability and acoustic signals: a multi-level approach in songbirds 
Biology Letters  2012;8(6):928-931.
Among songbirds, growing evidence suggests that acoustic adaptation of song traits occurs in response to habitat features. Despite extensive study, most research supporting acoustic adaptation has only considered acoustic traits averaged for species or populations, overlooking intraindividual variation of song traits, which may facilitate effective communication in heterogeneous and variable environments. Fewer studies have explicitly incorporated sexual selection, which, if strong, may favour variation across environments. Here, we evaluate the prevalence of acoustic adaptation among 44 species of songbirds by determining how environmental variability and sexual selection intensity are associated with song variability (intraindividual and intraspecific) and short-term song complexity. We show that variability in precipitation can explain short-term song complexity among taxonomically diverse songbirds, and that precipitation seasonality and the intensity of sexual selection are related to intraindividual song variation. Our results link song complexity to environmental variability, something previously found for mockingbirds (Family Mimidae). Perhaps more importantly, our results illustrate that individual variation in song traits may be shaped by both environmental variability and strength of sexual selection.
doi:10.1098/rsbl.2012.0522
PMCID: PMC3497114  PMID: 22859557
acoustic adaptation; birdsong; environmental variability; selection; sensory drive; song variation
10.  RUNAWAY SEXUAL SELECTION WITHOUT GENETIC CORRELATIONS: SOCIAL ENVIRONMENTS AND FLEXIBLE MATE CHOICE INITIATE AND ENHANCE THE FISHER PROCESS 
Female mating preferences are often flexible, reflecting the social environment in which they are expressed. Associated indirect genetic effects (IGEs) can affect the rate and direction of evolutionary change, but sexual selection models do not capture these dynamics. We incorporate IGEs into quantitative genetic models to explore how variation in social environments and mate choice flexibility influence Fisherian sexual selection. The importance of IGEs is that runaway sexual selection can occur in the absence of a genetic correlation between male traits and female preferences. Social influences can facilitate the initiation of the runaway process and increase the rate of trait elaboration. Incorporating costs to choice do not alter the main findings. Our model provides testable predictions: (1) genetic covariances between male traits and female preferences may not exist, (2) social flexibility in female choice will be common in populations experiencing strong sexual selection, (3) variation in social environments should be associated with rapid sexual trait divergence, and (4) secondary sexual traits will be more elaborate than previously predicted. Allowing feedback from the social environment resolves discrepancies between theoretical predictions and empirical data, such as why indirect selection on female preferences, theoretically weak, might be sufficient for preferences to become elaborated.
doi:10.1111/j.1558-5646.2012.01647.x
PMCID: PMC3627302  PMID: 22946795
Fisherian runaway; indirect genetic effects; interacting phenotypes; mate choice plasticity; mate preference learning; social evolution
11.  Sexual responsiveness is condition-dependent in female guppies, but preference functions are not 
BMC Ecology  2004;4:5.
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.
doi:10.1186/1472-6785-4-5
PMCID: PMC411045  PMID: 15117410
12.  Good Genes and Sexual Selection in Dung Beetles (Onthophagus taurus): Genetic Variance in Egg-to-Adult and Adult Viability 
PLoS ONE  2011;6(1):e16233.
Whether species exhibit significant heritable variation in fitness is central for sexual selection. According to good genes models there must be genetic variation in males leading to variation in offspring fitness if females are to obtain genetic benefits from exercising mate preferences, or by mating multiply. However, sexual selection based on genetic benefits is controversial, and there is limited unambiguous support for the notion that choosy or polyandrous females can increase the chances of producing offspring with high viability. Here we examine the levels of additive genetic variance in two fitness components in the dung beetle Onthophagus taurus. We found significant sire effects on egg-to-adult viability and on son, but not daughter, survival to sexual maturity, as well as moderate coefficients of additive variance in these traits. Moreover, we do not find evidence for sexual antagonism influencing genetic variation for fitness. Our results are consistent with good genes sexual selection, and suggest that both pre- and postcopulatory mate choice, and male competition could provide indirect benefits to females.
doi:10.1371/journal.pone.0016233
PMCID: PMC3022759  PMID: 21267411
13.  Intralocus sexual conflict for fitness: sexually antagonistic alleles for testosterone 
Intralocus sexual conflict occurs when a trait encoded by the same genetic locus in the two sexes has different optima in males and females. Such conflict is widespread across taxa, however, the shared phenotypic traits that mediate the conflict are largely unknown. We examined whether the sex hormone, testosterone (T), that controls sexual differentiation, contributes to sexually antagonistic fitness variation in the bank vole, Myodes glareolus. We compared (opposite-sex) sibling reproductive fitness in the bank vole after creating divergent selection lines for T. This study shows that selection for T was differentially associated with son versus daughter reproductive success, causing a negative correlation in fitness between full siblings. Our results demonstrate the presence of intralocus sexual conflict for fitness in this small mammal and that sexually antagonistic selection is acting on T. We also found a negative correlation in fitness between parents and their opposite-sex progeny (e.g. father–daughter), highlighting a dilemma for females, as the indirect genetic benefits of selecting reproductively successful males (high T) are lost with daughters. We discuss mechanisms that may mitigate this disparity between progeny quality.
doi:10.1098/rspb.2011.2340
PMCID: PMC3311893  PMID: 22171083
sex-limited epistasis; mammal; artificial selection; sexually antagonistic selection; sexual selection
14.  Socially flexible female choice differs among populations of the Pacific field cricket: geographical variation in the interaction coefficient psi (Ψ) 
Indirect genetic effects (IGEs) occur when genes expressed in one individual affect the phenotype of a conspecific. Theoretical models indicate that the evolutionary consequences of IGEs critically depend on the genetic architecture of interacting traits, and on the strength and direction of phenotypic effects arising from social interactions, which can be quantified by the interaction coefficient Ψ. In the context of sexually selected traits, strong positive Ψ tends to exaggerate evolutionary change, whereas negative Ψ impedes sexual trait elaboration. Despite its theoretical importance, whether and how Ψ varies among geographically distinct populations is unknown. Such information is necessary to evaluate the potential for IGEs to contribute to divergence among isolated or semi-isolated populations. Here, we report substantial variation in Ψ for a behavioural trait involved in sexual selection in the field cricket Teleogryllus oceanicus: female choosiness. Both the strength and direction of Ψ varied among geographically isolated populations. Ψ also changed over time. In a contemporary population of crickets from Kauai, experience of male song increased female choosiness. In contrast, experience of male song decreased choosiness in an ancestral population from the same location. This rapid change corroborates studies examining the evolvability of Ψ and demonstrates how interpopulation variation in the interaction coefficient might influence sexual selection and accelerate divergence of traits influenced by IGEs that contribute to reproductive isolation in nascent species or subspecies.
doi:10.1098/rspb.2012.0631
PMCID: PMC3396897  PMID: 22648156
indirect genetic effect; mate choice plasticity; psi; sexual selection; social environment
15.  Sex-specific effects of developmental environment on reproductive trait expression in Drosophila melanogaster 
Ecology and Evolution  2012;2(7):1362-1370.
Variation in the expression of reproductive traits provides the raw material upon which sexual selection can act. It is therefore important to understand how key factors such as environmental variation influence the expression of reproductive traits, as these will have a fundamental effect on the evolution of mating systems. It is also important to consider the effects of environmental variation upon reproductive traits in both sexes and to make comparisons with the environment to which the organism is adapted. In this study, we addressed these issues in a systematic study of the effect of a key environmental factor, variation in larval density, on reproductive trait expression in male and female Drosophila melanogaster. To do this, we compared reproductive trait expression when flies were reared under controlled conditions at eight different larval densities that covered a 20-fold range. Then, to place these results in a relevant context, we compared the results to those from flies sourced directly from stock cages. Many reproductive traits were surprisingly insensitive to variation in larval density. A notable exception was nonlinear variation in female fecundity. In contrast, we found much bigger differences in comparisons with flies from stock cages—including differences in body size, latency to mate, copulation duration, fecundity, and male share of paternity in a competitive environment. For a number of traits, even densities of 1000 larvae per vial (125 larvae per mL of food) did not phenocopy stock cage individuals. This study reveals novel patterns of sex-specific sensitivity to environmental variation that will influence the strength of sexual selection. It also illustrates the importance of comparisons with the environment to which individuals are adapted.
doi:10.1002/ece3.243
PMCID: PMC3434946  PMID: 22957145
Drosophila melanogaster; environmental variation; genotype-by-environment interaction; larval density; larval development; sexual selection
16.  Intraspecific sexual selection on a speciation trait, male coloration, in the Lake Victoria cichlid Pundamilia nyererei. 
The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciation. Extensive intra- and interspecific variation in male nuptial coloration and female mating preferences, in the absence of postzygotic isolation between species, has inspired the hypothesis that sexual selection has been a driving force in the origin of this species flock. This hypothesis rests on the premise that the phenotypic traits that underlie behavioural reproductive isolation between sister species diverged under sexual selection within a species. We test this premise in a Lake Victoria cichlid, by using laboratory experiments and field observations. We report that a male colour trait, which has previously been shown to be important for behavioural reproductive isolation between this species and a close relative, is under directional sexual selection by female mate choice within this species. This is consistent with the hypothesis that female choice has driven the divergence in male coloration between the two species. We also find that male territoriality is vital for male reproductive success and that multiple mating by females is common.
doi:10.1098/rspb.2004.2911
PMCID: PMC1691879  PMID: 15590594
17.  Both male and female identity influence variation in male signalling effort 
Background
Male sexual displays play an important role in sexual selection by affecting reproductive success. However, for such displays to be useful for female mate choice, courtship should vary more among than within individual males. In this regard, a potentially important source of within male variation is adjustment of male courtship effort in response to female traits. Accordingly, we set out to dissect sources of variation in male courtship effort in a fish, the desert goby (Chlamydogobius eremius). We did so by designing an experiment that allowed simultaneous estimation of within and between male variation in courtship, while also assessing the importance of the males and females as sources of courtship variation.
Results
Although males adjusted their courtship depending on the identity of the female (a potentially important source of within-male variation), among-male differences were considerably greater. In addition, male courtship effort towards a pair of females was highly repeatable over a short time frame.
Conclusion
Despite the plasticity in male courtship effort, courtship displays had the potential to reliably convey information about the male to mate-searching females. Our experiment therefore underscores the importance of addressing the different sources contributing to variation in the expression of sexually-selected traits.
doi:10.1186/1471-2148-11-233
PMCID: PMC3163561  PMID: 21827657
18.  Intrasexual competition in females: evidence for sexual selection? 
Behavioral Ecology  2011;22(6):1131-1140.
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.
doi:10.1093/beheco/arr106
PMCID: PMC3199163  PMID: 22479137
aggression; female competition; intrasexual selection; mating success; sexual selection
19.  Sexual selection and natural selection in bird speciation 
The role of sexual selection in speciation is investigated, addressing two main issues. First, how do sexually selected traits become species recognition traits? Theory and empirical evidence suggest that female preferences often do not evolve as a correlated response to evolution of male traits. This implies that, contrary to runaway (Fisherian) models of sexual selection, premating isolation will not arise as an automatic side effect of divergence between populations in sexually selected traits. I evaluate premating isolating mechanisms in one group, the birds. In this group premating isolation is often a consequence of sexual imprinting, whereby young birds learn features of their parents and use these features in mate choice. Song, morphology and plumage are known recognition cues. I conclude that perhaps the main role for sexual selection in speciation is in generating differences between populations in traits. Sexual imprinting then leads to these traits being used as species recognition mechanisms. The second issue addressed in this paper is the role of sexual selection in adaptive radiation, again concentrating on birds. Ecological differences between species include large differences in size, which may in themselves be sufficient for species recognition, and differences in habitat, which seem to evolve frequently and at all stages of an adaptive radiation. Differences in habitat often cause song and plumage patterns to evolve as a result of sexual selection for efficient communication. Therefore sexual selection is likely to have an important role in generating premating isolating mechanisms throughout an adaptive radiation. It is also possible that sexual selection, by creating more allopatric species, creates more opportunity for ecological divergence to occur. The limited available evidence does not support this idea. A role for sexual selection in accelerating ecological diversification has yet to be demonstrated.
doi:10.1098/rstb.1998.0207
PMCID: PMC1692216
20.  Coevolution of non-fertile sperm and female receptivity in a butterfly 
Biology Letters  2009;5(5):678-681.
Sexual conflict can promote rapid evolution of male and female reproductive traits. Males of many polyandrous butterflies transfer nutrients at mating that enhances female fecundity, but generates sexual conflict over female remating due to sperm competition. Butterflies produce both normal fertilizing sperm and large numbers of non-fertile sperm. In the green-veined white butterfly, Pieris napi, non-fertile sperm fill the females' sperm storage organ, switching off receptivity and thereby reducing female remating. There is genetic variation in the number of non-fertile sperm stored, which directly relates to the female's refractory period. There is also genetic variation in males' sperm production. Here, we show that females' refractory period and males' sperm production are genetically correlated using quantitative genetic and selection experiments. Thus selection on male manipulation may increase the frequency of susceptible females to such manipulations as a correlated response and vice versa.
doi:10.1098/rsbl.2009.0452
PMCID: PMC2781977  PMID: 19640869
sexual conflict; sperm competition; coevolution
21.  Genotype-by-Environment Interactions for Female Mate Choice of Male Cuticular Hydrocarbons in Drosophila simulans 
PLoS ONE  2013;8(6):e67623.
Recent research has highlighted the potential importance of environmental and genotype-by-environment (G×E) variation in sexual selection, but most studies have focussed on the expression of male sexual traits. Consequently, our understanding of genetic variation for plasticity in female mate choice is extremely poor. In this study we examine the genetics of female mate choice in Drosophila simulans using isolines reared across two post-eclosion temperatures. There was evidence for G×Es in female choosiness and preference, which suggests that the evolution of female mate choice behaviour could differ across environments. However, the ranked order of preferred males was consistent across females and environments, so the same males are favoured by mate choice in spite of G×Es. Our study highlights the importance of taking cross-environment perspectives in order to gain a more comprehensive understanding of the operation of sexual selection.
doi:10.1371/journal.pone.0067623
PMCID: PMC3692475  PMID: 23825675
22.  Age before beauty? Relationships between fertilization success and age-dependent ornaments in barn swallows 
Behavioral Ecology and Sociobiology  2011;65(9):1687-1697.
When males become more ornamented and reproduce more successfully as they grow older, phenotypic correlations between ornament exaggeration and reproductive success can be confounded with age effects in cross-sectional studies, and thus say relatively little about sexual selection on these traits. This is exemplified here in a correlative study of male fertilization success in a large colony of American barn swallows (Hirundo rustica erythrogaster). Previous studies of this species have indicated that two sexually dimorphic traits, tail length and ventral plumage coloration, are positively correlated with male fertilization success, and a mechanism of sexual selection by female choice has been invoked. However, these studies did not control for potential age-related variation in trait expression. Here, we show that male fertilization success was positively correlated with male tail length but not with plumage coloration. We also show that 1-year-old males had shorter tails and lower fertilization success than older males. This age effect accounted for much of the covariance between tail length and fertilization success. Still, there was a positive relationship between tail length and fertilization success among older males. But as this group consisted of males from different age classes, an age effect may be hidden in this relationship as well. Our data also revealed a longitudinal increase in both tail length and fertilization success for individual males. We argue that age-dependent ornament expression and reproductive performance in males complicate inferences about female preferences and sexual selection.
doi:10.1007/s00265-011-1176-4
PMCID: PMC3156913  PMID: 21949464
Extrapair paternity; Hirundo rustica; Life-history variation; Plumage coloration; Sexual selection; Tail length
23.  The immunocompetence handicap hypothesis: testing the genetic predictions 
The immunocompetence handicap hypothesis suggests that the immune system competes for resources with sexually selected ornaments; variation in ornaments might reflect genetic variation for immunocompetence. We tested this genetic prediction by mating scorpionfly females to males differing in the expression of a condition-dependent ornament trait, saliva secretion, and then comparing offspring immunocompetence. We found several indications of an immunocompetence handicap in our study: females had superior immunocompetence compared with males, the different immune traits were positively correlated, and there were indications of genetic variation in immune traits. However, we found no significant difference in the immunocompetence of offspring derived from males differing in ornament expression, only a tendency for sons of ornamented males to possess slightly better immunocompetence. The estimated effect of fathers on offspring immunocompetence was rather small, but it might be a sufficient benefit of female choice, provided that the costs of choice are small. We conclude that the genetic benefit of female choice is small concerning offspring immunocompetence, but the immunocompetence handicap principle might nevertheless work in scorpionflies.
PMCID: PMC1690479  PMID: 10693823
24.  Fin-mutant female zebrafish (Danio rerio) exhibit differences in association preferences for male fin length 
Behavioural processes  2008;80(1):35-38.
Females often choose to associate with males that have exaggerated traits. In fishes, this may reflect an overall preference for larger size in a potential mate. Female zebrafish (Danio rerio) prefer males with larger bodies but not longer fins. The availability of mutant and transgenic strains of zebrafish make this a unique model system in which to study the role of phenotypic variation in social and sexual behavior. We used mutant strains of zebrafish with truncated (short fin) and exaggerated (long fin) fins to further examine female preferences for fin length in dichotomous association tests. Wild type females showed no preferences between wild type males and short fin mutant males or between wild type males and long fin mutant males. short fin females also showed no preference for short fin males or wild type males while long fin females preferred to associate with long fin males over wild type males. These results suggest that the single gene long fin mutation that results in altered fin morphological may also be involved in a related female association preference.
doi:10.1016/j.beproc.2008.09.004
PMCID: PMC3418694  PMID: 18848866
association preference; Danio rerio; long fin; short fin; zebrafish
25.  Female Influence on Pre- and Post-copulatory Sexual Selection and its Genetic Basis in Drosophila melanogaster 
Molecular ecology  2011;20(19):4098-4108.
Genetic variation among females is likely to influence the outcome of both pre- and post-copulatory sexual selection in Drosophila melanogaster. Here we use association testing to survey natural variation in 10 candidate female genes for their effects on female reproduction. Females from 91 chromosome 2 substitution lines were scored for phenotypes affecting pre- and post-copulatory sexual selection such as mating and remating rate, propensity to use sperm from the second male to mate, and measures of fertility. There were significant genetic contributions to phenotypic variation for all the traits measured. Resequencing of the 10 candidate genes in the 91 lines yielded 68 nonsynonymous polymorphisms which were tested for associations with the measured phenotypes. Twelve significant associations (markerwise P < 0.01) were identified. Polymorphisms in the putative serine protease homolog CG9897 and the putative odorant binding protein CG11797 associated with female propensity to remate and met an experimentwise significance of P < 0.05. Several other associations, including those impacting both fertility and female remating rate suggest that sperm storage might be an important factor mitigating female influence on sexual selection.
doi:10.1111/j.1365-294X.2011.05253.x
PMCID: PMC3183161  PMID: 21902747
Sperm competition; association testing; female mating; sexual selection; genotype-phenotype

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