Evolutionary dynamics of a sexual ornament in the house sparrow (Passer domesticus): the role of indirect selection within and between sexes

The relative contribution of sexual and natural selection to evolution of sexual ornaments has rarely been quantified under natural conditions. In this study we used a long-term dataset of house sparrows in which parents and offspring were matched genetically to estimate the within- and across-sex genetic basis for variation and covariation among morphological traits. By applying two-sex multivariate "animal models" to estimate genetic parameters, we estimated evolutionary changes in a male sexual ornament, badge size, from the contribution of direct and indirect selection on correlated traits within males and females, after accounting for overlapping generations and age-structure. Indirect natural selection on genetically correlated traits in males and females was the major force causing evolutionary change in the male ornament. Thus, natural selection on female morphology may cause indirect evolutionary changes in male ornaments. We observed however no directional phenotypic change in the ornament size of one-year-old males during the study period. On the other hand, changes were recorded in other morphological characters of both sexes. Our analyses of evolutionary dynamics in sexual characters require application of appropriate two-sex models to account for how selection on correlated traits in both sexes affects the evolutionary outcome of sexual selection.     

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.     

On the origins of sexual dimorphism in butterflies

The processes governing the evolution of sexual dimorphism provided a foundation for sexual selection theory. Two alternative processes, originally proposed by Darwin and Wallace, differ primarily in the timing of events creating the dimorphism. In the process advocated by Darwin, a novel ornament arises in a single sex, with no temporal separation in the origin and sex-limitation of the novel trait. By contrast, Wallace proposed a process where novel ornaments appear simultaneously in both sexes, but are then converted into sex-limited expression by natural selection acting against showy coloration in one sex. Here, we investigate these alternative modes of sexual dimorphism evolution in a phylogenetic framework and demonstrate that both processes contribute to dimorphic wing patterns in the butterfly genera Bicyclus and Junonia. In some lineages, eyespots and bands arise in a single sex, whereas in other lineages they appear in both sexes but are then lost in one of the sexes. In addition, lineages displaying sexual dimorphism were more likely to become sexually monomorphic than they were to remain dimorphic. This derived monomorphism was either owing to a loss of the ornament ('drab monomorphism') or owing to a gain of the same ornament by the opposite sex ('mutual ornamentation'). Our results demonstrate the necessity of a plurality in theories explaining the evolution of sexual dimorphism within and across taxa. The origins and evolutionary fate of sexual dimorphism are probably influenced by underlying genetic architecture responsible for sex-limited expression and the degree of intralocus sexual conflict. Future comparative and developmental work on sexual dimorphism within and among taxa will provide a better understanding of the biases and constraints governing the evolution of animal sexual dimorphism.     

Signal trait sexual dimorphism and mutual sexual selection in Drosophila serrata

Abstract The evolution of sexual dimorphism may occur when natural and sexual selection result in different optimum trait values for males and females. Perhaps the most prominent examples of sexual dimorphism occur in sexually selected traits, for which males usually display exaggerated trait levels, while females may show reduced expression of the trait. In some species, females also exhibit secondary sexual traits that may either be a consequence of a correlated response to sexual selection on males or direct sexual selection for female secondary sexual traits. In this experiment, we simultaneously measure the intersex genetic correlations and the relative strength of sexual selection on males and females for a set of cuticular hydrocarbons in Drosophila serrata . There was significant directional sexual selection on both male and female cuticular hydrocarbons: the strength of sexual selection did not differ among the sexes but males and females preferred different cuticular hydrocarbons. In contrast with many previous studies of sexual dimorphism, intersex genetic correlations were low. The evolution of sexual dimorphism in D. serrata appears to have been achieved by sex-limited expression of traits controlled by genes on the X chromosome and is likely to be in its final stages.     

Sexual selection in the barn swallow (Hirundo rustica). V. Geographic variation in ornament size

Models of sexual selection in a cline predict the patterns of clinal variation in female mate preference and male secondary sexual characters. These predictions were tested for the nominate subspecies of the barn swallow Hirundo rustica which demonstrates clinal variation in morphology, with several characters in both sexes showing increasing size at higher latitudes. Sexual size dimorphism in the length of the tail ornament and the short, central tail feathers increase with increasing latitude while size dimorphism in other morphological characters is independent of latitude. The main reason for the two divergent patterns of sexual size dimorphism appears to be the higher foraging cost of having a long tail ornamental at low latitudes. The control of development decreases with increasing latitude as demonstrated by an increasing latitudinal cline in fluctuating asymmetry of tail length. Phenotypic variance in tail length increases with latitude in males, but not in females, as shown by the coefficients of variation. Clinal variation in morphology is not due to natural selection associated with a latitudinal increase in the distance between breeding and wintering areas. The geographic patterns of morphological variation suggest that the tail character has diverged geographically as a result of a sexual process of reliable signalling.     

Patterns of fluctuating asymmetry in sexual ornaments predict female choice

Extravagant secondary sexual characters are assumed to have arisen and be maintained by sexual selection. While traits like horns, antlers and spurs can be ascribed to intrasexual competition, other traits such as extravagant feather ornaments, displays and pheromones have to be ascribed to mate choice. A number of studies have tested whether females exert selection on the size of male ornaments, but only some of these have recorded female preferences for the most extravagantly ornamented males. Here I demonstrate that female choice can be directly predicted from the relationship between the degree of fluctuating asymmetry and the size of a secondary sexual character. Fluctuating asymmetry is an epigenetic measure of the ability of individuals to cope with stress, and it occurs when an individual is unable to undergo identical development of an otherwise bilaterally symmetric trait on both sides of its body. There is a negative relationship between the degree of fluctuating asymmetry and the absolute size of an ornament in those bird species with a female preference for the largest male sex trait, while there is a flat or U-shaped relationship among species without a female preference. These results suggest that females prefer exaggerated secondary sexual characters if they reliably demonstrate the ability of males to cope with genetic and environmental stress. Some species may demonstrate a flat or U-shaped relationship between the degree of fluctuating asymmetry and the absolute size of an ornament because (i) the genetic variance in viability signalled by the secondary sex trait has been depleted; (ii) the secondary sex trait is not particularly costly and therefore does not demonstrate condition dependence; or because (iii) the sex traits can be considered arbitrary traits rather than characters reflecting good genes.     

Sexual selection, natural selection and the evolution of dimorphic coloration and ornamentation in agamid lizards

Both sexual selection and natural selection can influence the form of dimorphism in secondary sexual traits. Here, we used a comparative approach to examine the relative roles of sexual selection and natural selection in the evolution of sexually dimorphic coloration (dichromatism) and ornamentation in agamid lizards. Sexual dimorphism in head and body size were used as indirect indicators of sexual selection, and habitat type (openness) as an index of natural selection. We examined separately the dichromatism of body regions "exposed to" and "concealed from" visual predators, because these body regions are likely to be subject to different selection pressures. Dichromatism of "exposed" body regions was significantly associated with habitat type: males were typically more conspicuously coloured than females in closed habitats. By contrast, dichromatism of "concealed" body regions and ornament dimorphism were positively associated with sexual size dimorphism (SSD). When we examined male and female ornamentation separately, however, both were positively associated with habitat openness in addition to snout-vent length and head SSD. These results suggest that natural selection constrains the evolution of elaborate ornamentation in both sexes as well as sexual dichromatism of body regions exposed to visual predators. By contrast, dichromatism of "concealed" body regions and degree of ornament dimorphism appear to be driven to a greater degree by sexual selection.     

No fecundity cost of female secondary sexual trait expression in the horned beetle Onthophagus sagittarius

Typically males bear the products of sexual selection in the form of ornaments and/or weapons used to compete for and attract females. Secondary sexual traits in females have been thought of as the product of correlated responses to sexual selection on males. However, there is increasing phylogenetic evidence that female secondary sexual traits can arise independently of selection on males, and may be subject to sexual selection. Theoretical models of the evolution of female ornamentation via male mate choice have assumed that females suffer a cost of ornament expression via reduced fecundity, and hence female ornaments are less likely to evolve than male ornaments. In the dung beetle Onthophagus sagittarius, there has been an independent evolutionary origin of horns in females that are qualitatively different from the horns produced by males. We use this system as a model to examine the costs of horn expression for females within a life-history context. We identified a longevity cost of reproduction for females that was independent of horn expression. Large females lived longer, and after controlling for lifespan, had a higher lifetime fecundity, and invested more heavily in maternal provisioning than did small females. We found no evidence of a cost to females of investment in horns. Rather, the rate of increase in fecundity and horn expression with body size were equal, so that absolute horn size provides an accurate indicator of body size and maternal quality. The effects we observe were independent of female contest competition and/or male mate choice, which were excluded in our experimental protocol. However, we speculate on the potential functional contributions female horns might make to female fitness.     

Sexual dimorphism in wing beat frequency in relation to eye span in stalk‐eyed flies (Diopsidae)

Although male ornaments may provide benefits to individuals bearing them, such structures may also entail fitness costs. Selection should favour aspects of the phenotype that act to reduce such costs, yet such compensatory traits are often ignored in studies of sexual selection. If a male ornament increases predation risk via reduced locomotor performance, then there may be selection for changes in morphological traits to compensate for behavioural or biomechanical changes in how individuals use their morphology (or both). We took a comparative approach aiming to test whether changes in wing beat frequency are evolutionarily correlated with increases in male ornamentation across stalk‐eyed fly species. Previous studies have shown that increased male eye span is evolutionarily correlated with increased wing size; thus, we tested whether there is additional compensation via increases in size‐adjusted wing beat frequency. The results obtained revealed that relative wing beat frequency is negatively related to relative eye span in males, and sexual dimorphism in wing beat frequency is negatively related to dimorphism in eye span. These findings, in addition to our finding that eye span dimorphism is positively related to aspect ratio dimorphism, suggest that male stalk‐eyed flies compensate primarily by increasing wing size and shape, which may then have resulted in the subsequent evolutionary reduction in wing beat frequency. Thus, exaggerated ornaments can result in evolutionary modifications in wing morphology, which in turn lead to adjustments in flapping kinematics, illustrating the tight envelope of trade‐offs when compensating for exaggerated ornaments. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 670–679.     

Timing of ornament growth, phenotypic variation, and size dimorphism in two promiscuous African whydahs (Ploceidae: Vidua)

Variation within populations is a prerequisite for the action of selection on morphological traits. Darwin assumed that there was much greater variation in sexual ornament size than in body size, but this may not be generally true of natural populations. I analyse field data on variation in body size and the length, area and mass of tail ornaments in paradise (Vidua paradisaea) and shaft-tailed whydahs (V. regia). Whydahs are promiscuous, brood parasitic African finches with elaborate tail ornaments in breeding males. The short, unadorned tails of male shaft-tailed whydahs, which carry a wire-like tail ornament, are non-significantly (1%) longer than female tails, but male paradise whydahs, which carry a large, broad ornament, have unadorned tails 10% longer than those of conspecific females. Fully grown ornament length, mass and area vary little more (CVs = 1.8-6.4%) than male or female body size traits (CVs = 1.7-6.1%). Instead, there is high variation in the timing of ornament development during prenuptial moult (CVs = 30.8–39.5% for paradise whydahs and 12.6–23.8% for shaft-tailed whydahs when corrected to a standard date). This temporal variation in development probably has greater significance for sexual selection in whydahs than maximum ornament size.     

Haematocrit correlates with tail ornament size in three populations of the Barn Swallow (Hirundo rustica)

1. Handicap models of sexual selection propose that male ornaments are indicators of male quality and that honesty is enforced by the costs imposed by the exaggerated ornamental traits. In long-distance migratory birds that feed on the wing, the aerodynamic cost of exaggerated ornamental characters should be particularly high because the size of the ornaments deviates from the natural selection optimum. During migration, birds are expected to raise their oxygen consumption in relation to the energetic demands imposed by their morphology. An increase of haematocrit is an adaptive response to enhance oxygen uptake and efficiency of transfer to the muscular tissues during spells of intense muscular activity.
2. The change of haematocrit of Barn Swallows ( Hirundo rustica ) after their arrival to the breeding sites, and the relationships between haematocrit values recorded after migration and the size of ordinary and sexually selected morphological characters in three Barn Swallow populations were analysed.
3. Males had higher haematocrit values than females. Individual haematocrit values declined after arrival to the breeding sites. Haematocrit values of males were significantly and positively correlated with the size of their ornamental tail but not correlated with other characters, thus suggesting that well-ornamented males, in order to arrive early, have to raise their haematocrit above the level of short-tailed males.
4. Males and females of similar tail length did not differ in their haematocrit, thus suggesting that sexual dimorphism in haematocrit might be functionally related to dimorphism in tail length.
5. Our results are consistent with the handicap principle because long-tailed males experience lower mortality and larger seasonal reproductive success compared with short-tailed males.     

Testing the interactive effects of testosterone and parasites on carotenoid‐based ornamentation in a wild bird

Abstract Testosterone underlies the expression of most secondary sexual traits, playing a key role in sexual selection. However, high levels might be associated with physiological costs, such as immunosuppression. Immunostimulant carotenoids underpin the expression of many red‐yellow ornaments, but are regulated by testosterone and constrained by parasites. We manipulated testosterone and nematode burdens in red grouse (Lagopus lagopus scoticus) in two populations to tease apart their effects on carotenoid levels, ornament size and colouration in three time‐step periods. We found no evidence for interactive effects of testosterone and parasites on ornament size and colouration. We showed that ornament colouration was testosterone‐driven. However, parasites decreased comb size with a time delay and testosterone increased carotenoid levels in one of the populations. This suggests that environmental context plays a key role in determining how individuals resolve the trade‐off between allocating carotenoids for ornamental coloration or for self‐maintenance needs. Our study advocates that adequately testing the mechanisms behind the production or maintenance of secondary sexual characters has to take into account the dynamics of sexual trait expression and their environmental context.     

Humoral immune response in relation to senescence, sex and sexual ornamentation in the barn swallow (Hirundo rustica)

Performance of animals may decline with age. The effects of senescence, however, may differ between the sexes because of differences in physiology and behaviour. Acquired immunity provides hosts with efficient mechanisms of anti-parasite defence, but the effect of senescence on immunocompetence has never been studied in natural populations. In the barn swallow (Hirundo rustica), primary antibody response to an antigen during one breeding season declined with age in females, while secondary response during the following breeding season declined with age in both sexes. Parasite-mediated sexual selection theory posits that male secondary sexual characters reveal resistance to parasites. Males with large tail ornaments had stronger primary response, retained larger antibody levels until the following year, but did not differ in secondary response compared with short-tailed males, as predicted if ornamentation reflects resistance to parasites. This is the first study showing that immunocompetence declines with age in any vertebrate under natural conditions.     

HOW TO COMPENSATE FOR COSTLY SEXUALLY SELECTED TAILS: THE ORIGIN OF SEXUALLY DIMORPHIC WINGS IN LONG-TAILED BIRDS

Recent work on birds suggests that certain morphological differences between the sexes may have evolved as an indirect consequence of sexual selection because they offset the cost of bearing extravagant ornaments used for fighting or mate attraction. For example, long-tailed male sunbirds and widowbirds also have longer wings than females, perhaps to compensate for the aerodynamic costs of tail elaboration. We used comparative data from 57 species to investigate whether this link between sexual dimorphism in wing and tail length is widespread among long-tailed birds. We found that within long-tailed families, variation in the extent of tail dimorphism was associated with corresponding variation in wing dimorphism. One nonfunctional explanation of this result is simply that the growth of wings and tails is controlled by a common developmental mechanism, such that long-tailed individuals inevitably grow long wings as well. However, this hypothesis cannot account for a second pattern in our data set: as predicted by aerodynamic theory, we found that, comparing across long-tailed families, sexual dimorphism in wing length varied with tail shape as well as with sex differences in tail length. Thus, wing dimorphism was generally greater in species with aerodynamically costly graduated tails than in birds with cheaper, streamer-shaped tails. This result was not caused by confounding phylogenetic effects, because it persisted when phylogeny was controlled for, using an independent comparisons method. Our findings therefore confirm that certain aspects of sexual dimorphism may sometimes have evolved through selection for traits that reduce the costs of elaborate sexually selected characters. We suggest that future work aimed at understanding sexual selection by investigating patterns of sexual dimorphism should attempt to differentiate between the direct and indirect consequences of sexual selection.     

The allometric pattern of sexually size dimorphic feather ornaments and factors affecting allometry

The static allometry of secondary sexual characters is currently subject to debate. While some studies suggest an almost universal positive allometry for such traits, but isometry or negative allometry for nonornamental traits, other studies maintain that any kind of allometric pattern is possible. Therefore, we investigated the allometry of sexually size dimorphic feather ornaments in 67 species of birds. We also studied the allometry of female feathers homologous to male ornaments (female ornaments in the following) and ordinary nonsexual traits. Allometries were estimated as reduced major axis slopes of trait length on tarsus length. Ornamental feathers showed positive allometric slopes in both sexes, although that was not a peculiarity for ornamental feathers, because nonsexual tail feathers also showed positive allometry. Migration distance (in males) and relative size of the tail ornament (in females) tended to be negatively related to the allometric slope of tail feather ornaments, although these results were not conclusive. Finally, we found an association between mating system and allometry of tail feather ornaments, with species with more intense sexual selection showing a smaller degree of allometry of tail ornaments. This study is consistent with theoretical models that predict no specific kind of allometric pattern for sexual and nonsexual characters.     

Variable sexual ornaments in scarlet-tufted malachite sunbirds (Nectarinia johnstoni) on Mount Kenya

In order to be elaborated by sexual selection, sexual ornaments must vary perceptibly and genetically among individuals in natural populations. Rather little is known about ornament variation in monogamous species, in which sexual selection should act more weakly than in polygynous species. We report phenotypic variation in feather ornament size (elongated tails and pectoral tufts) and body size in the scarlet-tufted malachite sunbird Nectarinia johnstoni , a monogamous, sexually dimorphic nectarivore of East African alpine zones. Fully-expressed male ornaments are highly significantly more variable (CVs = 12–29%) than are skeletal and wing measures primarily affected by natural selection (CVs = 2 4%). Female sunbirds have pectoral tufts which are significantly (22–25%) smaller than those of adult males, but more variable (CV_s= 21–22%, CV_s= 12–15%), and more variable than body size. Among males with fully-grown ornaments, those with longer tails tend to have longer wings and wider tufts. The high variation in fully-grown ornaments in malachite sunbirds is consistent with the view that the ornaments are condition-dependent sexual signals. Finally, we review studies of feather ornament variation to date, and show that ornaments are much more variable in monogamous than non-monogamous species, apparently due to the relatively weak pressure of sexual selection.     

Aerodynamic costs of long tails in male barn swallows Hirundo rustica and the evolution of sexual size dimorphism

Exaggerated tail feathers of birds constitute a standard exampleof evolution of extravagant characters due to sexual selection.Such secondary sexual traits are assumed to be costly to produceand maintain, and they usually are accompanied by morphologicaladaptations that tend to reduce their costs. The aerodynamiccosts for male barn swallows Hirundo rustica of having longtails were quantified using aerodynamics theory applied to morphologicaldata from seven European populations. Latitudinal differencesin tail length were positively correlated with differences inflight costs predicted by aerodynamics theory. A positive relationshipbetween aerodynamic costs of long tails and the degree of sexualsize dimorphism was found among populations. Latitudinal differencesin foraging costs may result in tail length being relativelysimilar in males and females in southern populations, whereasthe low foraging costs for males in northern populations mayallow them to cope with higher aerodynamic costs, giving riseto large sexual size dimorphism. Enlargement of wingspan inmales can alleviate but not eliminate the costs of tail exaggeration,and therefore differences in aerodynamic costs of male ornamentswere maintained among populations. Sexual size dimorphism in thebarn swallow arises as a consequence of latitudinal differencesin the advantages of sexual selection for males and the costsof long tails for males and females.     

Offspring sexual dimorphism and sex-allocation in relation to parental age and paternal ornamentation in the barn swallow

We analysed the morphology of nestling barn swallows (Hirundo rustica) in relation to their sex, and laying and hatching order. In addition, we studied sex-allocation in relation to parentage, parental age and expression of a secondary sexual character of fathers. Molecular sexing was conducted using the sex chromosome-linked avian CHD1 gene. Sex of the offspring was not associated with laying or hatching order. None of nine morphological, serological and immunological variables varied in relation to offspring sex. Sexual dimorphism did not vary in relation to parental age and expression of a paternal secondary sexual character. The proportion of sons declined with brood size. Individual males and females had a similar proportion of sons during consecutive breeding years. The proportion of sons of individual females declined with age, but increased with the expression of a secondary sexual character of their current mate. The generalized lack of variation in sexual dimorphism among nestlings may suggest that barn swallows do not differentially invest in sons vs. daughters. Alternatively, male offspring may require different parental effort compared to their female siblings in order to attain the same morphological state. The lack of variation in offspring sexual dimorphism with paternal ornamentation suggests no adjustment of overall parental effort in relation to reproductive value of the two sexes. However, male-biased sex ratio among offspring of highly ornamented males may represent an adaptive sex-allocation strategy because the expression of male ornaments is heritable and highly ornamented males are at a sexual selection advantage.     

BREEDING COMPETITION IN A PACIFIC SALMON (COHO: ONCORHYNCHUS KISUTCH): MEASURES OF NATURAL AND SEXUAL SELECTION

In the breeding system of Pacific salmon, females compete for oviposition territories, and males compete to fertilize eggs. The natural selection in females and sexual selection in males likely has been responsible for their elaborate breeding morphologies and the dimorphism between the sexes. We quantified direct-selection intensities during breeding on mature coho salmon (Oncorhynchus kisutch), measured for seven phenotypic characters, including three secondary sexual characters. Wild and sea-ranched hatchery coho were used to enhance the range of phenotypes over which selection could be examined. The fish were allowed to breed in experimental arenas where we could quantify components of breeding success as well as estimate overall breeding success. We found that without competition, natural selection acts only on female body size for increased egg production; there is no detectable selection on males for the phenotypic distribution we used. Under competition, the opportunity for selection increased sixfold among females. Natural selection favored female body size and caudal-peduncle (tail) depth. Increased body size meant increased egg production and access to nesting territories. The caudal peduncle, used in burst swimming and nest digging, influenced both successful egg deposition and nest survival. Increasing density increased competition among females, though it did not significantly intensify natural selection on their characters. In males, competition increased the opportunity for selection 52-fold, which was nine times greater than for females. Sexual selection favored male body size and hooked snout length, both characters directly influencing male access to spawning opportunities. Selection on male body size was also affected significantly by breeding density. The ability of large males to control access to spawning females decreased at higher densities reflecting an increase in the operational sex ratio. Further, the relative success of small males, which could sneak access to spawning females, appeared to increase as that of intermediate-sized males decreased. Such disruptive selection may be responsible for the evolution of alternative reproductive tactics in salmon.     

No evidence of assortative mating on the basis of putative ornamental traits in Long‐tailed Finches Poephila acuticauda

When multiple ornaments are expressed in both sexes, they are generally assumed to be maintained by mutual sexual selection and have a function in mate choice. In the Long‐tailed Finch Poephila acuticauda both sexes exhibit multiple ornaments that vary in their expression in either size (pintail and throat patch) or colour (bill) between individuals and sexes. We assessed whether these ornaments are maintained by mutual sexual selection by exploring whether individuals in a wild population paired assortatively with respect to these ornamental traits, and the degree to which the expression of these ornamental traits was indicative of reproductive success. We found no evidence of assortative pairing with respect to variation in homologous ornaments or body condition in the two sexes. In addition, we found no effect of ornament expression on the reproductive success of either males or females. Our findings suggest that the expression of these apparently ornamental traits in both sexes of this species may play no current role in mutual mate selection or as indicator traits of reproductive performance. We are currently unable to identify any function for these very elaborate ornaments in either sex of this species and suggest that the typical assumption that all such traits have an ornamental function may need further examination.