Selection on multiple sexual signals in two Central and Eastern European populations of the barn swallow

Variation in intensity and targets of sexual selection on multiple traits has been suggested to play a major role in promoting phenotypic differentiation between populations, although the divergence in selection may depend on year, local conditions or age. In this study, we quantified sexual selection for two putative sexual signals across two Central and East European barn swallow (Hirundo rustica rustica) populations from Czech Republic and Romania over multiple years. We then related these differences in selection to variation in sexual characters among barn swallow populations. Our results show that tail length and ventral coloration vary between populations, sexes, and age classes (first‐time breeders vs. experienced birds). We found that selection on tail length was stronger in first‐time breeders than in experienced birds and in males than in females in the Romanian population, while these differences between age groups and sexes were weak in Czech birds. We suggest that the populational difference in selection on tail length might be related to the differences in breeding conditions. Our results show that ventral coloration is darker (i.e., has lower brightness) in the Romanian than in the Czech population, and in experienced birds and males compared with first‐time breeders and females, respectively. The sexual difference in ventral coloration may suggest sexual selection on this trait, which is supported by the significant directional selection of ventral coloration in first‐time breeding males on laying date. However, after controlling for the confounding effect of wing length and tarsus length, the partial directional selection gradient on this trait turned nonsignificant, suggesting that the advantage of dark ventral coloration in early breeding birds is determined by the correlated traits of body size. These findings show that ventral coloration may be advantageous over the breeding season, but the underlying mechanism of this relationship is not clarified.     

Female butterflies prefer males bearing bright iridescent ornamentation

Butterflies are among nature's most colorful animals, and provide a living showcase for how extremely bright, chromatic and iridescent coloration can be generated by complex optical mechanisms. The gross characteristics of male butterfly colour patterns are understood to function for species and/or sex recognition, but it is not known whether female mate choice promotes visual exaggeration of this coloration. Here I show that females of the sexually dichromatic species Hypolimnas bolina prefer conspecific males that possess bright iridescent blue/ultraviolet dorsal ornamentation. In separate field and enclosure experiments, using both dramatic and graded wing colour manipulations, I demonstrate that a moderate qualitative reduction in signal brightness and chromaticity has the same consequences as removing the signal entirely. These findings validate a long-held hypothesis, and argue for the importance of intra- versus interspecific selection as the driving force behind the exaggeration of bright, iridescent butterfly colour patterns.     

Genetic basis and fitness correlates of dynamic carotenoid‐based ornamental coloration in male and female common kestrels Falco tinnunculus

Knowledge of the genetic basis of sexual ornaments is essential to understand their evolution through sexual selection. Although carotenoid‐based ornaments have been instrumental in the study of sexual selection, given the inability of animals to synthesize carotenoids de novo, they are generally assumed to be influenced solely by environmental variation. However, very few studies have directly estimated the role of genes and the environment in shaping variation in carotenoid‐based traits. Using long‐term individual‐based data, we here explore the evolutionary potential of a dynamic, carotenoid‐based ornament (namely skin coloration), in male and female common kestrels. We first estimate the amount of genetic variation underlying variation in hue, chroma and brightness. After correcting for sex differences, the chroma of the orange‐yellow eye ring coloration was significantly heritable (h² ± SE = 0.40 ± 0.17), whereas neither hue (h² = 0) nor brightness (h² = 0.02) was heritable. Second, we estimate the strength and shape of selection acting upon chromatic (hue and chroma) and achromatic (brightness) variation and show positive and negative directional selection on female but not male chroma and hue, respectively, whereas brightness was unrelated to fitness in both sexes. This suggests that different components of carotenoid‐based signals traits may show different evolutionary dynamics. Overall, we show that carotenoid‐based coloration is a complex and multifaceted trait. If we are to gain a better understanding of the processes responsible for the generation and maintenance of variation in carotenoid‐based coloration, these complexities need to be taken into account.     

Elevational plumage divergence in the Rufous-capped Babbler (Cyanoderma ruficeps) on a mountainous island

Environment plays an important role in the evolution of plumage coloration in birds and may also lead to sexual dichromatism if males and females face different selection pressures. Mountains exhibit varying ecological conditions along their elevation gradient that may impose divergent selection on elevationally widespread species, causing intraspecific plumage divergence. For example, UV light environments often vary between montane and lowland habitats, which could potentially cause differences in plumage UV reflection between birds occurring in the two types of habitats. However, few studies have examined the effects of elevation on plumage evolution. In this study, we quantified the plumage coloration of the Rufous-capped Babbler Cyanoderma ruficeps from montane and lowland habitats on a mountainous island, Taiwan. We aimed to examine whether their plumage showed differences associated with changing ecological environments across the elevational gradient. The results supported that the plumage of babblers occupying montane habitats had higher UV-reflectance and brightness than that of lowland birds, corresponding to the higher UV intensity in montane than lowland background light environments. The elevational differences were mainly found across the ventral parts of babblers that had relatively higher levels of UV reflectance compared with their dorsal parts. Alternatively, the brighter plumage, with higher UV-reflectance in montane than lowland birds, might be mediated by physiological adaptation to other ecological factors, such as parasite pressures. The elevational differences in plumage UV-reflectance and brightness were more dramatic in males than in females. However, we found significant sexual dichromatism in different body parts between montane and lowland babblers in which females had brighter or stronger UV-associated coloration than males, suggesting that sexual selection has little impact on babbler plumage. Our study suggests the importance of elevational divergent selection associated with UV light or other ecological environments on avian plumage evolution.     

Effects of polyandry on male phenotypic diversity

Polyandry has the potential to affect the distribution of phenotypes and to shape the direction of sexual selection. Here, we explore this potential using Trinidadian guppies as a model system and ask whether polyandry leads to directional and/or diversifying selection of male phenotypic traits. In this study, we compare the phenotypic diversity of offspring from multiply and singly sired broods. To quantify phenotypic diversity, we first combine phenotypic traits using multivariate methods, and then take the dispersion of individuals in multivariate space as our measure of diversity. We show that, when each trait is examined separately, polyandry generates offspring with a higher proportion of bright coloration, indicating directional selection. However, our multivariate approach reveals that this directionality is accompanied by an increase in phenotypic diversity. These results suggest that polyandry (i) selects for the production of sons with the preferred brighter colour phenotypes whereas (ii) enhancing the diversity of male sexual traits. Promoting phenotypic diversity may be advantageous in coping with environmental and reproductive variability by increasing long‐term fitness.     

Condition dependence, quantitative genetics, and the potential signal content of iridescent ultraviolet butterfly coloration

Structural colors result from an interaction between light and the fine-scale physical structure of a surface, and are often extremely bright, chromatic, and iridescent. Given that these visual features depend upon the aggregate abundance and architectural precision of photonic structures, structurally colored sexual ornaments seem well placed to indicate a range of mate quality characteristics. We tested this hypothesis by investigating the signaling potential of structural coloration in the sexually dimorphic butterfly Colias eurytheme. Males of this species display iridescent ultraviolet (UV) markings (arising from multilayer thin films) that overlay a broad area of yellowish-orange pigmentation on their dorsal wing surface. Only the structural UV has demonstrated function as a sexual signal; hence we predicted that it should contain more reliable phenotypic and/or genetic quality information, which would be indicated by phenotypic and/or genetically mediated condition dependence. In two split-family breeding experiments we manipulated condition by exposing full siblings to different stressors at two different juvenile life-history stages: (1) reduced larval host-plant quality and (2) transient heat/cold shocks during metamorphosis. Both stressors had profound effects on key developmental and life-history traits. Each stressor also significantly affected male dorsal coloration; thus, the expression of both structural and pigmentary coloration is phenotypically condition dependent. As predicted, the strongest condition dependence was evident in the brightness and angular visibility (i.e., iridescence) of the UV. Characteristics of both the iridescent UV and pigmentary orange also exhibited moderate-high and significant heritability (H(2) approximately h(2) approximately 0.4-0.9). However, genetic and residual variances did not increase under stress; thus, the observed condition dependence was not genetically mediated as predicted if wing color trait signals "good" genes for the ability to either withstand or circumvent developmental stress. The heightened stress sensitivity of the iridescent UV suggests that it offers an informative lifetime indicator of juvenile environments and, henceforth, adult male phenotypic condition, which may be salient to females seeking a highly fertile and/or nutritious male ejaculate.     

Conspicuous Female Ornamentation and Tests of Male Mate Preference in Threespine Sticklebacks (Gasterosteus aculeatus)

Sexual selection drives the evolution of exaggerated male ornaments in many animal species. Female ornamentation is now acknowledged also to be common but is generally less well understood. One example is the recently documented red female throat coloration in some threespine stickleback (Gasterosteus aculeatus) populations. Although female sticklebacks often exhibit a preference for red male throat coloration, the possibility of sexual selection on female coloration has been little studied. Using sequential and simultaneous mate choice trials, we examined male mate preferences for female throat color, as well as pelvic spine color and standard length, using wild-captured threespine sticklebacks from the Little Campbell River, British Columbia. In a multivariate analysis, we found no evidence for a population-level mate preference in males, suggesting the absence of directional sexual selection on these traits arising from male mate choice. Significant variation was detected among males in their preference functions, but this appeared to arise from differences in their mean responsiveness across mating trials and not from variation in the strength (i.e., slope) of their preference, suggesting the absence of individual-level preferences as well. When presented with conspecific intruder males, male response decreased as intruder red throat coloration increased, suggesting that males can discriminate color and other aspects of phenotype in our experiment and that males may use these traits in intrasexual interactions. The results presented here are the first to explicitly address male preference for female throat color in threespine sticklebacks.     

Selection and inheritance of sexually dimorphic juvenile plumage coloration

Sexually dimorphic plumage coloration is widespread in birds and is generally thought to be a result of sexual selection for more ornamented males. Although many studies find an association between coloration and fitness related traits, few of these simultaneously examine selection and inheritance. Theory predicts that sex‐linked genetic variation can facilitate the evolution of dimorphism, and some empirical work supports this, but we still know very little about the extent of sex linkage of sexually dimorphic traits. We used a longitudinal study on juvenile Florida scrub‐jays (Aphelocoma coerulescens) to estimate strength of selection and autosomal and Z‐linked heritability of mean brightness, UV chroma, and hue. Although plumage coloration signals dominance in juveniles, there was no indication that plumage coloration was related to whether or not an individual bred or its lifetime reproductive success. While mean brightness and UV chroma are moderately heritable, hue is not. There was no evidence for sex‐linked inheritance of any trait with most of the variation explained by maternal effects. The genetic correlation between the sexes was high and not significantly different from unity. These results indicate that evolution of sexual dimorphism in this species is constrained by low sex‐linked heritability and high intersexual genetic correlation.     

Signal design and courtship presentation coincide for highly biased delivery of an iridescent butterfly mating signal

Sensory drive theory contends that signaling systems should evolve to optimize transmission between senders and intended receivers, while minimizing visibility to eavesdroppers where possible. In visual communication systems, the high directionality afforded by iridescent coloration presents underappreciated avenues for mediating this trade-off. This hypothesis predicts functional links between signal design and presentation such that visual conspicuousness is maximized only under ecologically relevant settings and/or to select audiences. We addressed this prediction using Hypolimnas bolina, a butterfly in which males possess ultraviolet markings on their dorsal wing surfaces with a narrow angular reflectance function. Males bearing brighter dorsal markings are increasingly attractive to females, but also likely more conspicuous to predators. Our data indicate that, during courtship (and given the ritualized wingbeat dynamics at these times), males position themselves relative to females in such a way as to simultaneously maximize three components of known or putative signal conspicuousness: brightness, area, and iridescent flash. This suggests that male signal design and display have coevolved for the delivery of an optimally conspicuous signal to courted females. More broadly, these findings imply a potential signaling role for iridescence itself, and pose a novel example for how signal design may coevolve with the behavioral context of display.     

THE EVOLUTION OF SEXUAL DIMORPHISM BY SEXUAL SELECTION: THE SEPARATE EFFECTS OF INTRASEXUAL SELECTION AND INTERSEXUAL SELECTION

Libellula luctuosa, a pond dragonfly found in eastern North America, is apparently sexually dimorphic. Previous studies of the mating behavior in this species suggested that both male-male competition and female mate choice are important influences. Males compete for territories, where they attract females and where mating occurs. Female behavior influences both the copulation success and the fertilization success of males. Because of temporal and spatial separation of these episodes of sexual selection, multivariate and nonparametric statistical techniques could be used to investigate the influence of components of sexual selection on various sexually dimorphic traits. Sexual dimorphism in L. luctuosa was first quantified; then the direct effects and the form of selection were estimated. Sexually dimorphic wing size, body size, wing coloration, and body coloration are distributed either continuously or discontinuously between the sexes in L. luctuosa. These traits have apparently diverged between the sexes as a result of directional sexual selection. Body size is further influenced by stabilizing selection. Intrasexual selection (success in gaining access to a territory) and intersexual selection (success in copulation and fertilization) can influence the same or different sexually dimorphic characters. Body size is influenced by directional selection during the intrasexual phase of sexual selection and is also influenced by stabilizing selection during intersexual selection. The size of the brown wing patch is influenced by directional selection, primarily during the intersexual phase of sexual selection. There is directional selection on the white wing patch during both phases. Thus, the different proximate mechanisms of sexual selection may jointly or separately affect the evolution of sexually dimorphic characters. Further empirical and theoretical investigations into the differences in the effects of intrasexual selection and intersexual selection are needed to clarify the circumstances leading to separate consequences of these two mechanisms of sexual selection.     

Fecundity selection does not vary along a large geographical cline of trait means in a passerine bird

Local environmental and ecological conditions are commonly expected to result in local adaptation, although there are few examples of variation in phenotypic selection across continent‐wide spatial scales. We collected standardized data on selection with respect to the highly variable plumage coloration of pied flycatcher (Ficedula hypoleuca Pall.) males from 17 populations across the species' breeding range. The observed selection on multiple male coloration traits via the annual number of fledged young was generally relatively weak. The main aim of the present study, however, was to examine whether the current directional selection estimates are associated with distance to the sympatric area with the collared flycatcher (Ficedula albicollis Temminck), a sister species with which the pied flycatcher is showing character displacement. This pattern was expected because plumage traits in male pied flycatchers are changing with the distance to these areas of sympatry. However, we did not find such a pattern in current selection on coloration. There were no associations between current directional selection on ornamentation and latitude or longitude either. Interestingly, current selection on coloration traits was not associated with the observed mean plumage traits of the populations. Thus, there do not appear to be geographical gradients in current directional fecundity selection on male plumage ornamentation. The results of the present study do not support the idea that constant patterns in directional fecundity selection would play a major role in the maintenance of coloration among populations in this species. By contrast, the tendency for relatively weak mosaic‐like variation in selection among populations could reflect just a snapshot of temporally variable, potentially environment‐dependent, selection, as suggested by other studies in this system. Such fine‐grained variable selection coupled with gene flow could maintain extensive phenotypic variation across populations. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 808–827.     

Reversed plumage ontogeny in a female hummingbird: implications for the evolution of iridescent colours and sexual dichromatism

In polygynous birds, bright plumage is typically more extensive in the sexually competitive males and develops at or after sexual maturity. These patterns, coupled with the importance of male plumage in sexual displays, fostered the traditional hypothesis that bright plumages and sexual dichromatism develop through the actions of sexual selection on males. This view remains problematic for hummingbirds, all of which are polygynous, because their bright iridescent plumages are also important non-sexual signals associated with dominance at floral nectar sources. Here I show that female amethyst-throated sunangels [ Heliangelus amethysticollis (d'Orbigny & Lafresnaye)], moult from an immature plumage with an iridescent gorget to an adult plumage with a non-iridescent gorget. This 'reversed' ontogeny contradicts the notion that iridescent plumage has a sexual function because sexual selection in polygynous birds should be lowest among non-reproductive immature females. Moreover, loss of iridescent plumage in adult females indicates that adult sexual dichromatism in H. amethysticollis is due in large part to changes in female ontogeny. I suggest that both the ontogeny and sexual dichromatism evolved in response to competition for nectar.     

Rapid divergence of social signal coloration across the White Sands ecotone for three lizard species under strong natural selection

Animal social signals are important for population recognition, communication, and mate choice. Although natural selection often favours cryptic coloration, sexual selection can underlie patterns of coloration that function in inter- or intrasexual communication. We compared social signal coloration of three lizard species across a substrate colour ecotone in New Mexico. These species exhibit cryptic blanched dorsal coloration on the gypsum dunes of White Sands and dark coloration on the surrounding desert soils. We detected corresponding population divergence in colour used for intra- ( Aspidoscelis inornata , Sceloporus undulatus ) or inter- ( Holbrookia maculata ) sexual signalling. Although the magnitude and direction of change in coloration varied among taxa, differences in hue and chroma accounted for more variation in social coloration than for dorsal coloration. The relative conspicuousness of social signals also varied across the ecotone. We discuss the possibilities that divergent signalling colours in this system are the result of: (1) stochastic processes, (2) direct selection, and/or (3) a correlated response to natural selection on dorsal coloration.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 243–255.     

Interacting selection diversifies warning signals in a polytypic frog: an examination with the strawberry poison frog

Aposematic signals represent one of the most accessible traits to evaluate the interaction of natural and sexual selection on signal evolution. Here we investigate the contributions of these two selective forces on the aposematic signal evolution of the highly polytypic strawberry poison frog, Oophaga pumilio, of Bocas del Toro, Panama. Previous research has shown that the brightness of O. pumilio warning coloration can inform predators of the toxicity levels associated with different populations of the archipelago. Other studies suggest that sexual selection may be influencing warning signal brightness within populations via female mate choice (Isla Solarte, Isla Bastimentos, and Aquacate Peninsula populations) and male–male competition (Isla Solarte). Here we present two non-exclusive scenarios for how natural and sexual selection interact to drive phenotypic variation across this archipelago: (1) predators impose a selective regime whereby populations above a toxicity-brightness threshold are at liberty to diversify via sexual selection and below which populations are constrained to maintain a stricter resemblance to a more cryptic population mean, and (2) synergistic/additive effects of inter- and intrasexual selection drive the evolution of brighter males within populations above this toxicity threshold. We investigate whether aposematic patterns of divergence across the archipelago relative to the common mainland phenotype meet these predictions using existing data on O. pumilio morph toxicity measures and overall conspicuousness estimates to an avian predator. Using standardized z-scores to evaluate the range of trait values, we find that indeed the population representative of the common mainland phenotype (Almirante) represents an intermediate level of both toxicity and conspicuousness, and that derived Bocas del Toro populations vary in each of those components in directions predicted by the proposed scenarios. Furthermore, we find greater divergence towards conspicuousness than crypsis, a pattern suggestive of sexual and natural selection acting synergistically in morphs with high toxicity.     

Contrasting patterns of selection on the size and coloration of a female plumage ornament in common yellowthroats

Females often possess ornaments that appear smaller and duller than homologous traits in males. These ornaments may arise as nonfunctional by‐products of sexual selection in males and cause negative viability or fecundity selection in females in proportion to the cost of their production and maintenance. Alternatively, female ornaments may function as signals of quality that are maintained by sexual or social selection. In a 4‐year study of 83 female common yellowthroats (Geothlypis trichas) and their 222 young, we found strong viability and fecundity selection on the yellow bib, a carotenoid‐based plumage ornament that is a target of sexual selection in males. Females with larger bibs were older, larger and more fecund than females with smaller bibs. However, bib size positively covaried with bib total brightness and carotenoid chroma, aspects of bib coloration that were under negative viability and fecundity selection. Females with more colourful bibs laid fewer eggs in their first clutch, were more likely to suffer total brood loss due to predation and were less likely to return to the study area. Selection against bib coloration limits the value of bib size as a quality indicator in females and may constrain the elaboration of bib attributes in males.     

Comparing plumage colour measurements obtained directly from live birds and from collected feathers: the case of the great tit Parus major

Birds frequently display a colourful plumage which is important both in inter and intraespecific communication, and either in sexual and social contexts. In last years some methodologies have been developed to, analyse plumage coloration, but the use of the spectrometers has been particularly important for UV range. Measurement of plumage coloration with the spectrometer may be taken directly on the bird or, alternatively by collecting some feathers and measuring them later in the laboratory. However, few is known about the reliability of measures obtained from feathers and whether these are really representative of plumage coloration. We tested this assumption analysing measurements of carotenoids-based coloration components (lightness, chroma and hue) and lutein peak of the yellow breast of the great tit Parus major. We used two spectrometers (Ocean optics and Minolta) which calculate differently the colour components. Our results showed that direct measurement of bird was highly repeatable to determine lightness, chroma and hue for both spectrometers. Similar results we found for collected feathers procedure for both devices. Collected feathers provided high representative measurements of colour values with Minolta spectrometer. Lightness was highly repeatable when we used Ocean optic spectrometer, but chroma and hue were moderate. Lutein peak was also highly repeatable in all cases. The number of feathers used to measure plumage coloration in collected feathers procedure strongly influenced values of colour plumage variables. In general, values of lightness, chroma and hue stabilised when more than 10–15 feathers were used although we found slight differences between spectrometers. However, only four feathers were needed for lutein peak. Thus, our results stress the need to use a minimum number of feathers in measuring plumage coloration from collected feathers.     

SEXUAL SELECTION AND SURVIVAL SELECTION ON WING COLORATION AND BODY SIZE IN THE RUBYSPOT DAMSELFLY HETAERINA AMERICANA

I review methodological problems that can lead to false evidence for selection on secondary sexual characters and present a study of selection in rubyspot damselflies (Hetaerina americana) that avoids these pitfalls. Male rubyspots have a large red spot on each wing that grows to a terminal size after sexual maturity. Selection gradient analyses revealed evidence for positive sexual and survival selection on both terminal wing spot size and body size. Phenotype manipulations confirmed that wing spot size was subject to direct sexual selection, but showed that the positive slope of survival on wing spot size was an indirect effect of selection on unmeasured traits. This study provides the strongest evidence yet for sexual selection on coloration in Odonata, but also provides clear examples of why phenotypic selection statistics must be calculated and interpreted cautiously.     

Reduced sexual dichromatism,mutual ornamentation,and individual quality in the monogamous Zenaida dove Zenaida aurita

Although variation in plumage coloration is known to occur both between and within sexes, its study remains limited to a few bird families. The Zenaida dove Zenaida aurita is a socially monogamous tropical columbid bird species, characterized by an overall cinnamon‐brownish plumage and structural colorations on the head and neck. The species has been described as sexually dichromatic for plumage, although color differences between males and females are not obvious in the field. We investigated variation in the presumably melanin‐based color of the crown, mantle, breast, and belly, in the iridescent dark‐blue streaks on the head, and in the symmetric iridescent patches on the neck, over the whole spectrum visible to birds. Further, unlike most previous studies, we assessed covariation between plumage color and phenotypic traits in both males and females in relation to the putative signaling function of ornaments. Zenaida doves appeared to be slightly sexually dichromatic for the hue of pigment‐based colored areas, with males being on average more reddish than females. However, this difference was not discernible when considering the avian visual system. Conversely, although the reflectance spectra of iridescent plumage did not significantly differ between sexes in brightness, chroma or spectral position of the peaks, color discrimination analyses showed that individuals should be able to perceive between‐ or within‐sex differences in the color of the iridescent patch. In addition, several color parameters of brown and iridescent feathers were significantly related to territorial status, body condition, wing chord, and, albeit weakly, to individual multilocus heterozygosity. Overall, our results thus suggest that plumage color might be a reliable signal of quality in individuals of both sexes in this species. Further studies are needed to test the potential implication of plumage coloration in mate choice and mating patterns in the Zenaida dove.     

Effects of cuticle structure and crystalline wax coverage on the coloration in young and old males of Calopteryx splendens and Calopteryx virgo

Male secondary sexual characters, such as color patterns, are often investigated at the macroscale level. However, micro- and nanoscale levels of morphological investigations may reveal functional features responsible for a particular coloration, thus providing more information, e.g., about the condition dependence of male sexual characters. The aim of this paper was to investigate cuticle color and its structure in males of two congeneric damselfly species, Calopteryx splendens and Calopteryx virgo, and reveal possible color changes with age. According to spectrometer measurements, C. splendens males were bluer and had a greater saturation of blue in their abdomen than C. virgo males, which were, in turn, greener and had more green saturation. Although the two species differed in the number of structural layers and the spacing of the layers, it seems that intactness of the wax crystals covering the epicuticle was most often the morphological trait which was related to the color parameters measured from males’ cuticles. The effect of the crystalline wax coverage on cuticle color was also confirmed by removing the wax using chloroform: after the treatment, the hue was bluer, the cuticle had a greater brightness and greater blue saturation, but less green saturation. Age differences influencing the color and structure of the cuticle were also observed: older males had more blue and green saturation and had more intact wax coverage than did younger males. Although multilayer reflection should be responsible for the iridescent color of males, our results suggest that wax coverage plays an important role in the color tuning of the male cuticle. This may have a considerable signal function, indicating the males’ viability to competing males or to females.     

Female mating biases for bright ultraviolet iridescence in the butterfly Eurema hecabe (Pieridae)

Exaggerated male-limited coloration is widespread among butterflies,yet convincing demonstrations of intraspecific mating preferencesfor signal brightness and/or chromaticity are relatively rarein this group. Here, I couple behavioral experiments involvingmanipulations of ambient light environments and male reflectancepatterns with observation of wild mating patterns to investigatevisual mating biases in the large grass yellow (Eurema hecabe).Males in this species possess exaggerated, limited-view ultraviolet(UV) iridescence across most of their dorsal wing surface thathas putative sexual signaling function. In the first experiment,conducted in small (0.7–m³) cages, individuals were significantlyless likely to copulate when the UV portion of natural ambientillumination (i.e., 300–400 nm) was strongly reduced.In 2 subsequent experiments, conducted under full-spectrum sunlightin small and large (5 x 6 x 4 m) cages, males with their UVsignal artificially dulled by 25% consistently copulated withfewer, and smaller, females than sham-control individuals. Importantly,the manipulated levels of UV brightness in these experimentsfall well within the naturally occurring bounds of variationin male UV reflectance. These findings therefore unanimouslysupport the presence of a UV signal–based female bias.In apparent contrast, comparison of 161 in-copula and 188 free-flyingmales from a high-density field assemblage revealed that copulatingmales were significantly older and henceforth actually possessed(subtly) less UV bright wings. Copulating male UV brightnesswas, however, positively related to the size of their mate,which echoes the experimental findings and may represent a signatureof mutual mate choice. I discuss these results in light of thefull complexities of the butterfly mating system and the potentialsignaling value of iridescent coloration in butterflies andanimals generally.