Dual functions of a melanin-based ornament in the common yellowthroat

Melanin-based ornaments often function as signals in male-male competition, whereas carotenoid-based ornaments appear to be important in female mate choice. This difference in function is thought to occur because carotenoid pigments are more costly to produce than melanins and are thus more reliable indicators of male quality. We examined the role of melanin- and carotenoid-based ornaments in male-male competition and female choice in the common yellowthroat Geothlypis trichas, a sexually dichromatic passerine. Males display a black facial mask produced by melanin pigmentation and a bright yellow bib (throat, breast and belly) produced by carotenoid pigmentation. In controlled aviary experiments, mask size was the best predictor of both male-male competition and female mate choice, and, therefore, mask size may be regarded as an ornament of dual function. These dual functions may help to maintain the reliability of mask size as an indicator of male quality, despite the potentially low cost of production. The size of the bib was unrelated to male-male competition or female choice, but there was a tendency for females to prefer males with more colourful bibs. We propose that the black mask is important in competition for territories with other males and for attracting females. Our results highlight the need for more studies of the mechanisms of sexual selection in species with ornaments composed of different pigment types.     

Female mate choice and male–male competition in Tonkean macaques: Who decides?

The theory of sexual selection predicts that females should be discriminatory in the choice of sexual partners. Females can express their choice in two ways. In direct mate choice, they show preferences for certain partners. In indirect mate choice, they select partners by displaying sexually attractive traits, thus eliciting contest competition between males. We focused on a primate species in which females advertise the timing of their ovulation and studied the balance between these two choice strategies. We tested predictions related to three hypotheses about direct and indirect female choice, namely the best‐male, graded‐signal and weak‐selectivity hypotheses. We investigated the sexual and agonistic interactions occurring during oestrous periods in five captive groups of Tonkean macaques (Macaca tonkeana). The results showed that dominant males used mate guarding to monopolise sexual access to parous females that were in the fertile stage of their reproductive cycle, while lower‐ranking males monitored only nulliparous females. The distribution of sexual presentations indicated that females accepted different types of partners, supporting the weak‐selectivity hypothesis regarding direct mate choice. The analysis of behavioural sequences revealed that mate‐guarding males used mild coercive behaviours to prevent females from mating with other males at conception time. The distribution of mounts showed that females mainly mated with dominant males, which leads us to argue that the best‐male hypothesis provides the most parsimonious explanation regarding indirect mate choice in Tonkean macaques. At the individual level, it may be concluded that male competitive strategies prevented females from exercising direct mate choice. At the evolutionary level, however, female sexual advertising and thus indirect choice promoted competition between males. The outcome is that indirect mate choice appears more important than direct mate choice in female Tonkean macaques.     

Preference for Male Traits Differ in Two Female Morphs of the Tree Lizard,Urosaurus ornatus

Non-random female mating preferences may contribute to the maintenance of phenotypic variation in color polymorphic species. However, the effect of female preference depends on the types of male traits used as signals by receptive females. If preference signals derive from discrete male traits (i.e., morph-specific), female preferences may rapidly fix to a morph. However, female preference signals may also include condition-dependent male traits. In this scenario, female preference may differ depending on the social context (i.e., male morph availability). Male tree lizards (Urosaurus ornatus) exhibit a dewlap color polymorphism that covaries with mating behavior. Blue morph males are aggressive and defend territories, yellow males are less aggressive and defend smaller territories, and orange males are typically nomadic. Female U. ornatus are also polymorphic in dewlap color, but the covariation between dewlap color and female behavior is unknown. We performed an experiment to determine how female mate choice depends on the visual and chemical signals produced by males. We also tested whether female morphs differ in their preferences for these signals. Female preferences involved both male dewlap color and size of the ventral color patch. However, the female morphs responded to these signals differently and depended on the choice between the types of male morphs. Our experiment revealed that females may be capable of distinguishing among the male morphs using chemical signals alone. Yellow females exhibit preferences based on both chemical and visual signals, which may be a strategy to avoid ultra-dominant males. In contrast, orange females may prefer dominant males. We conclude that female U. ornatus morphs differ in mating behavior. Our findings also provide evidence for a chemical polymorphism among male lizards in femoral pore secretions.     

Male pipefish prefer dominant over attractive females

Animals may obtain information guiding their choice betweenpotential partners from observing competitive interactionsand displays between them, or from displays directed at thechoosing individual. In the sex-role reversed pipefish Syngnathustyphle females display a temporary ornament (a color pattern)to other females as well as to males. We have previously shown that display of female ornaments per se is attractive to males.Here we show that information from competitive displays canoverride such direct attraction displays as signals in thepartner choice process. In a mate choice experiment, an enclosedmale could choose between two females. On the first experimentalday, females could interact freely, while on the second daythey were isolated from each other. When female-female competitionwas allowed, the ornament display was directed more to theother female than to the male: Time competing, rather thantime courting the male, correlated with ornament display duration.However, ornament display under competition and ornament displayin the absence of competition did not correlate significantly.In fact, females competing more intensively on day one displayedthe ornament less on day two. Furthermore, the ornament displayduring the first, but not the second, day predicted male matechoice on the second day. Thus, males remembered previous informationfrom competitive displays and used it rather than immediateinformation from displays in the absence of female-female competition.We suggest that competitive displays more reliably signal female quality as compared to noncompetitive ones, and that males benefitfrom mating with dominant females.     

Sex-role reversal revisited: choosy females and ornamented, competitive males in a pipefish

In the pipefish Syngnathus typhle sex roles are reversed, thatis, females compete more intensely than males over mates. However,competition over mates among individuals of one sex does notnecessarily prevent members of that same sex from being choosy,and choosiness in the other sex does not prevent competitionwithin it. In an experiment we allowed a female pipefish tochoose freely between two males, after which we released themales and let the three interact. Comparisons with earlier resultsshow that both sexes courted partners and competed with consexuals.However, females courted more often than did males, and courtshipwas more frequent in treatments involving large individualsthan in treatments with small individuals. Males competed amongthemselves for access to mates but for a shorter duration thanfemales in the same situation. Males displayed an ornament towardsfemales but not to males during mating competition. Females,however, used their ornament in both contexts. Females did notalways mate with the male of their previously made choice, whichwe interpret as females being constrained by male-male competition,male motivation to mate, or both. Thus, in this sex-role reversedspecies, mate choice in the more competitive sex may be circumventedand even overruled by mate competition and mating willingnessin the least competitive sex. Hence, sex roles should not beconsidered as sexes being either choosy or competitive but ratherthat males and females may exhibit different combinations ofchoice and competition.     

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.     

Female choice for sick males over healthy males: Consequences for offspring

Sexual selection theory indicates that ornament expression in males is in close relation to their condition. This “honesty” relationship serves as the basis for female choice: Females would mate with healthy males over sick males after assessing male ornament signal expression and derive benefits for their progeny. Here, we investigated female mate choice for infected and non‐infected males, male survival after infection (to corroborate the negative effect of infection), and fitness consequences of female preferences using Tenebrio molitor beetles. Male infection was produced having two types of challenges as follows: males infected with entomopathogenic fungi and males infected with nylon implants. Similar to previous studies, we corroborated that females preferred fungus‐infected males over positive control, negative control, and nylon‐challenged males. Survival was the lowest for fungus‐treated males followed by nylon‐treated and control males. Females mated with fungus‐treated males laid fewer and smaller eggs, and the laid eggs had less lipid content with a reduced eclosion success compared to females mated with non‐challenged males. Our interpretation is that fungus‐treated males invested their energetic resources to increase their attractiveness at the risk of survival, in a terminal investment fashion. Females, however, would have corrected their choice by investing less in their offspring.     

An intimidating ornament in a female pipefish

A sexually selected signal may serve a dual function being bothattractive to mates and deterring rivals. Presently, there arefew unambiguous demonstrations of an ornament functioning inboth a mate choice and mate competition context and none regardingfemale ornaments. We have shown earlier that a temporary ornament,a striped pattern, in a sex-role reversed female pipefish, Syngnathustyphle, attracts males. Here we show that this ornament alsointimidates rival females: in one experiment a male could interactwith either 1 or 2 females. Latency until copulation was longerwhen 2, rather than 1, females were present. Moreover, when2 females were present, competition lasted longer and time untilmating took place increased when females displayed their ornamentsmore equally. In another experiment, a focal female could see1 stimulus female and 1 stimulus male, the latter 2 being unawareof each other. The ornament of the stimulus female was manipulated,either strengthened by being painted black or left unalteredby being sham-painted. As a result, focal females experiencingblack-painted stimulus females decreased courtship as well ascompetitive activities compared with focal females seeing sham-paintedfemales. Moreover, focal females seeing black-painted femalesdisplayed less of their own ornament compared with controls.This decrease was due to a decrease in display toward malesrather than to stimulus females. Thus, this female ornamentindeed has a dual function, attracting mates and deterring rivals.In addition, the social costs invoked by this intimidating effecton rivals may help to maintain signal honesty.     

Ornament size and colour as alternative strategies for effective communication in gliding lizards

Sexual ornamentation needs to be conspicuous to be effective in attracting potential mates and defending territories and indeed, a multitude of ways exists to achieve this. Two principal mechanisms for increasing conspicuousness are to increase the ornament's colour or brightness contrast against the background and to increase the size of the ornament. We assessed the relationship between the colour and size of the dewlap, a large extendible throat‐fan, across a range of species of gliding lizards (Agamidae; genus Draco) from Malaysia and the Philippines. We found a negative relationship across species between colour contrast against the background and dewlap size in males, but not in females, suggesting that males of different species use increasing colour contrast and dewlap size as alternative strategies for effective communication. Male dewlap size also increases with increasing sexual size dimorphism, and dewlap colour and brightness contrast increase with increasing sexual dichromatism in colour and brightness, respectively, suggesting that sexual selection may act on both dewlap size and colour. We further found evidence that relative predation intensity, as measured from predator attacks on models placed in the field, may play a role in the choice of strategy (high chromatic contrast or large dewlap area) a species employs. More broadly, these results highlight that each component in a signal (such as colour or size) may be influenced by different selection pressures and that by assessing components individually, we can gain a greater understanding of the evolution of signal diversity.     

Ultraviolet nuptial colour determines fight success in male European green lizards (Lacerta viridis)

Animal communication through colour signals is a central theme in sexual selection. Structural colours can be just as costly and honest signals as pigment-based colours. Ultraviolet (UV) is a structural colour that can be important both in intrasexual competition and mate choice. However, it is still unknown if a UV signal alone can determine the outcome of male-male fights. European green lizard (Lacerta viridis) males develop a nuptial throat coloration with a strong UV component. Among males differing only in their manipulated UV colour, females prefer males with higher UV. Here, we experimentally decreased the UV coloration of randomly chosen males from otherwise similar male pairs to test the hypothesis that a difference in UV colour alone can affect fight success during male-male competition. Our results fully supported the hypotheses: in almost 90 per cent of the contests the male with reduced UV lost the fight. Our results show that UV can be an important signal, affecting both female mate choice and determining male fight success.     

Egg Colour Covaries with Female Expression of a Male Ornament in the Spotless Starling (Sturnus unicolor)

The sexually selected egg colour hypothesis (SSECH) proposes that egg colouration is as a post-mating sexually selected signal of female phenotypic quality, maintained by a higher allocation of paternal care. Similarly, some female traits can reflect genetic quality or condition and males could use this information in mate choice or in modulating parental investment. In our study, we examined the correlation of individual variation in egg colouration with female expression of a male ornament and how male feeding covaried with these two female traits in the spotless starling, in which egg colour varies widely between clutches and where both sexes possess showy throat feathers that are age dependent and that may signal individual quality. According to the SSECH, high-quality females (females with longer throat feathers) are expected to lay more colourful eggs than low-quality females and males should modify their feeding behaviour accordingly. By means of a principal component analysis, we found that most of the variation in egg colouration was due to brightness differences, and in a lower proportion to chromatic variation. Chromatic variation reflected a ultraviolet (UV) vs. greenness trade-off and was positively associated with throat feather length: females with larger throat feathers laid eggs with higher UV and lower green reflectance. However, egg brightness was not related to female feather length, as the SSECH would predict. Male feedings were positively related to female throat feather length and negatively related to chromatic variation, meaning that males contributed more to nests of females with long throat feathers who laid eggs with higher UV and lower green reflectance. In conclusion, our data provide mixed support for the SSECH: although egg chromatic variation was related to female expression of a male ornament and male parental care, we found no evidence that egg brightness was involved in these processes.     

Male mate choice in fishes

Selection generally favours male competition for females, and female mate choice of males. If, however, females vary in quality, and if males are limited in the maximum number of females with which they can mate, then selection should also favour male mate choice. We report on male mate choice in two species of fishes with different mating systems: the threespine stickleback, which has male parental care, and the coho salmon, which has female parental care. In both species, males allocated their mating effort in direct proportion to female quality.     

Competitive PCR reveals the complexity of postcopulatory sexual selection in Teleogryllus commodus

The outcome of mate choice depends on complex interactions between males and females both before and after copulation. Although the competition between males for access to mates and premating choice by females are relatively well understood, the nature of interactions between cryptic female choice and male sperm competition within the female reproductive tract is less clear. Understanding the complexity of postcopulatory sexual selection requires an understanding of how anatomy, physiology and behaviour mediate sperm transfer and storage within multiply mated females. Here we use a newly developed molecular technique to directly quantify mixed sperm stores in multiple mating females of the black field cricket, Teleogryllus commodus. In this species, female postcopulatory choice is easily observed and manipulated as females delay the removal of the spermatophore in favour of preferred males. Using twice‐mated females, we find that the proportion of sperm in the spermatheca attributed to the second male to mate with a female (S₂) increases linearly with the time of spermatophore attachment. Moreover, we show that the insemination success of a male increases with its attractiveness and decreases with the size of the female. The effect of male attractiveness in this context suggests a previously unknown episode of mate choice in this species that reinforces the sexual selection imposed by premating choice and conflicts with the outcome of postmating male harassment. Our results provide some of the clearest evidence yet for how sperm transfer and displacement in multiply mated females can lead directly to cryptic female choice, and that three distinct periods of sexual selection operate in black field crickets.     

On the function of female ornaments: male bluethroats prefer colourful females

Female ornaments in animals with conventional sex roles have traditionally been considered non-functional, being merely a genetically correlated response to selection for male ornamentation. Alternatively, female ornaments may be influenced by selection acting directly on the females, either through female–female competition or male choice. We tested the latter hypothesis in mate choice experiments with bluethroats (Luscinia s. svecica), a passerine bird in which females vary considerably in coloration of an ornamental throat patch. In outdoor aviaries placed in prime breeding habitat, males were allowed to choose between a colourful and a drab female. We found that males associated more with, and performed more sexual behaviours towards, colourful females. Female coloration was not age-related, but correlated significantly with body mass and tarsus length. Thus, we have demonstrated both a male preference for female ornamentation, and a relationship between ornament expression and female body size, which may be indicative of quality. Our results refute the correlated response hypothesis and support the hypothesis that female ornamentation is sexually selected.     

Male Ornamentation in a Sex‐Role Reversed Pipefish Corythoichthys haematopterus

Sexual selection theory predicts that mating competition in sex‐role reversed animals acts more strongly on females than males and consequently females are expected to develop secondary sexual traits. However, in a sex‐role reversed pipefish Corythoichthys haematopterus (family: Syngnathidae), only males develop an ornamental trait on the thorax, consisting of approx. 3–5 speckles alternated by lateral stripes of brilliant light blue and orange. To understand the function of this male ornament, we examined whether the presence of females affects the expression of this trait, and whether the expression of this trait depends on the male’s physical condition. Individual males were reared in a tank for a month in four different conditions: in high or low food supply and in the presence or absence of a female. After 1 mo, males in better condition expressed larger and deeper blue and yellow speckles, and males maintained with a female expressed larger and deeper blue speckles than solely reared males. These results indicated that the male ornament functions as a signal conveying information on the phenotypic quality of its holder and that females are potential receivers of this signal. Because C. haematopterus exhibits strict monogamy and competition for a mate occurs only among females, we concluded that the male ornament is not displayed in the context of mating competition but rather it is used as a cue for partner recognition to maintain pair bond.     

I'm sexy and I glow it: female ornamentation in a nocturnal capital breeder

In many species, males rely on sexual ornaments to attract females. Females, by contrast, rarely produce ornaments. The glow-worm (Lampyris noctiluca) is an exception where wingless females glow to attract males that fly in search of females. However, little is known about the factors that promote the evolution of female ornaments in a sexual selection context. Here, we investigated if the female ornament of the glow-worm is a signal of fecundity used in male mate choice. In support of this, we found brightness to correlate with female fecundity, and males to prefer brighter dummy females. Thus, the glow emitted by females is a reliable sexual signal of female fecundity. It is likely that male preference for the fecundity-indicating ornament has evolved because of large variation among females in fecundity, and because nocturnal males cannot directly assess female size and fecundity. These results indicate that female ornamentation may evolve in capital breeders (i.e. those in which stored resources are invested in reproduction) when females vary significantly in fecundity and this variation cannot be assessed directly by males.     

Female Choice, Female Reluctance to Mate and Sexual Selection on Body Size in the Dung Fly Sepsis cynipsea

We investigated the mechanisms of sexual selection in the common dung fly Sepsis cynipsea and how these affect selection on body size at the population level. Because of the presumed costs associated with mating, we predicted that there would be a decrease in the general reluctance of females to mate with any particular male at higher male densities at the mating site, a fresh cow pat, resulting in indirect female choice and a decrease in the strength of sexual selection. In contrast, classical direct female choice and male‐male competition should result in increased selection intensities because more opportunities for choice and competition exist at higher densities. Female reluctance to mate and female assessment of males are expressed in prominent female behaviour to repel mates in several insect species, including S. cynipsea. Laboratory pair‐wise choice experiments showed that large males were more likely to obtain copulations, which also ensued more promptly, suggesting female assessment of male quality (direct female choice). There was a basic influence of male activity but little further effect of male scramble competition on the outcome of mating. Another laboratory experiment showed a decrease in female shaking duration per male, associated with an asymptote in the shaking duration per female, as male density and harassment increased, but did not show the increase in mating frequency predicted by the female reluctance hypothesis. A study estimating sexual selection differentials in the field showed that directional selection for larger males was present overall and was negatively related to seasonally mediated variation in male density. Our study suggests that direct female choice in combination with indirect female choice (due to an interaction of female reluctance to mate and male persistence) is most consistent with the behavioural and selection patterns observed in S. cynipsea, but male effects cannot be definitively excluded.     

Year-round resource defence and the evolution of male and female song in suboscine birds: social armaments are mutual ornaments

The evolution of sexually monomorphic (i.e. mutual) ornamentation has attracted growing attention as a 'blind-spot' in evolutionary biology. The popular consensus is that female ornaments are subject to the same modes of sexual selection as males: intrasexual competition and mate choice. However, it remains unclear how these forces interact within and between sexes, or whether they fully capture selection on female traits. One possibility is that the 'armament-ornament' model - which proposes that traits used primarily in male-male contests are also co-opted by females as indicators of male quality - can be extended to explain signal evolution in both sexes. We examine this idea by testing the function of acoustic signals in two species of duetting antbirds. Behavioural observations and playback experiments suggest that male and female songs function primarily as armaments in competitive interactions. Removal experiments reveal that song is also a classic ornament used by unpaired males and females to advertise for mates. These results indicate that 'armament-ornament' processes may operate in reciprocal format, potentially explaining widespread mutual ornamentation in species with elevated intrasexual competition for resources. In addition, given that songs mediate competition between species outside the breeding season, our findings suggest that processes shaping monomorphic ornaments extend beyond the traditional definitions of sexual selection and are best understood in the broader framework of social selection.     

Female mate choice in relation to heterozygosity in Tribolium castaneum

Abstract Female mate choice, both before and after copulation, is pervasive among insect species. It is often hypothesized that females would preferentially mate with males that are genetically dissimilar to promote the genetic variability of the offspring. We used various strains of red flour beetle, Tribolium castaneum, and tested the effect of male and female genetic backgrounds on precopulatory and post‐copulatory female mate choice. Simultaneous mate choice experiments using previously well established pheromone assays did not detect female preference for males of different strains. Post‐copulatory female mate choice was examined through paternity analysis. Two parameters were used to measure post‐copulatory female mate choice, including male defence capacity (P₁, proportion of offspring sired by the first male when a female mated with two males consecutively) and offence capacity (P₂, proportion of offspring sired by the second of two males to mate with a female). When female and male beetle strains were same, defence capacity was significantly higher than when female and male strains were different. However, such a pattern was not observed for offence capacity. The results suggest that female precopulatory mate choice is not affected by genetic background, but the outcome of post‐copulatory processes depends on the genetic background of male and female beetles.     

FEMALES RECEIVE A LIFE-SPAN BENEFIT FROM MALE EJACULATES IN A FIELD CRICKET

Abstract.— Mating has been found to be costly for females of some species because of toxic products that males transfer to females in their seminal fluid. Such mating costs seem paradoxical, particularly for species in which females mate more frequently than is necessary to fertilize their eggs. Indeed, some studies suggest that females may benefit from mating more frequently. The effect of male ejaculates on female life span and lifetime fecundity was experimentally tested in the variable field cricket, Gryllus lineaticeps. In field crickets, females will mate repeatedly with a given male and mate with multiple males. Females that were experimentally mated either repeatedly or multiply lived more than 32% longer than singly mated females. In addition, multiply mated females produced 98% more eggs than singly mated females. Because females received only sperm and seminal fluid from males in the experimental matings, these life‐span and fecundity benefits may result from beneficial seminal fluid products that males transfer to females during mating. Mating benefits rather than mating costs may be common in many animals, particularly in species where female mate choice has a larger effect on male reproductive success than does the outcome of sperm competition.