Achromatic plumage reflectance predicts reproductive success in male black-capped chickadees

The size of achromatic (black, white, gray) plumage patchesserves as a male status signal in many species of birds, butvariation in the colors of these patches has received littleattention. We assessed the relation between achromatic plumagereflectance, dominance rank, body condition, and reproductivesuccess in male black-capped chickadees, Poecile atricapillus.We measured plumage reflectance for five body regions of 40male chickadees in late winter and monitored these males throughoutthe following breeding season to determine whether they survivedto breed, whether they successfully paired, whether their partnerlaid eggs, and both their apparent and realized reproductivesuccess. As expected from past studies, a male's dominance ranksignificantly predicted whether his partner laid eggs. However,only achromatic plumage reflectance significantly predictedother measures of male reproductive performance. Among maleswho fledged at least one offspring, both the brightness of whiteplumage regions and the UV-chroma of melanin-based plumage regionswere significant predictors of the proportion of within-pairyoung in their nests. When we consider all males we measured,assigning zero values to males who failed to sire any offspring,the UV-chroma of melanin-based plumage regions was a significantpredictor of realized reproductive success. Bib size was alsorelated to male realized reproductive success. Our findingssuggest that individual variation in achromatic plumage mayplay an important role in sexual signaling in chickadees.     

The relative importance of male tail length and nuptial plumage on social dominance and mate choice in the red-backed fairy-wren Malurus melanocephalus: evidence for the multiple receiver hypothesis

Understanding why males of many species exhibit two or more sexual ornaments depends upon identifying both the information conveyed and the intended receiver(s) for each signal. Here we focus on identifying the intended receivers for two sexual signals exhibited by male red-backed fairy-wrens Malurus melanocephalus , extent of nuptial plumage and tail length. In doing so we test the multiple receiver hypothesis, which predicts that each trait is directed toward a different type of receiver (e.g., males vs females). Male red-backed fairy-wrens in nuptial plumage exhibit reversed sexual dimorphism for tail length in the breeding season, when their tails are significantly shorter than those of females or males in eclipse plumage. Using both aviary-based experiments and indices of mate choice and social dominance from a natural population, we found that extent of nuptial plumage and age primarily affected female mate choice and that shorter tails were primarily associated with male:male dominance signaling. The field and aviary studies combined are consistent with the multiple receiver hypothesis, in that each trait appears to be directed primarily to a different set of receivers (plumage for females and tail length for males), though each trait may also signal information to the other set of receivers as well. We propose that sexual selection may favor shorter tail lengths in male red-backed fairy-wrens through social competition mechanisms.     

White Flank Spots Signal Feeding Dominance in Female Diamond Firetails,Stagonopleura guttata

Plumage colour can be used as an honest signal to convey health and status, which has traditionally been examined in the sexual selection context of choosy females and elaborate males. We use a model avian system to study the role of plumage coloration in a social context such as inter‐ and intrasexual competition over food resources. The diamond firetail (Stagonopleura guttata) is an endemic Australian finch: females have more white flank spots than males, and white spot number was correlated with cell‐mediated immune response in females. We use two experimental designs to test the role of white flank spots for feeding dominance and dominance discrimination in a group‐living bird. The results from two‐choice trials and from single‐arena trials showed that female ornamentation was consistently important in social food contests, and males consistently responded to female spot number. Females with higher spot number fed first, in trials with males and/or females. Also, females preferred to feed next to test birds with low spot number, but males showed no preference for feeding next to birds with few or many spots. Finally, latency to feed was predicted by spot number: both males and females had longer latency to feed if test birds had more spots than the focal birds. We conclude that female, but not male, ornamentation was important for inter‐ and intrasexual food competition. This is one of the very few studies to show that the same plumage ornament can have a different function between the sexes as a signal of social status. Moreover, this study shows that white plumage can be a signal of dominance.     

Brightness variability in the white badge of the eagle owl Bubo bubo

The application of modern spectrometry to the study of avian colour variability has revealed ignored patterns of colour variation such as male‐biased sexual dichromatism and seasonal variability in the plumage. However, the variation in the achromatic property of such traits, that is in the total light reflectance of the spectrum (i.e., brightness), has commonly been overlooked. The evolution of signals based on brightness should be favoured in those species that are active when light is scarce, i.e. at dawn and dusk. The eagle owl Bubo bubo is monogamous and apparently monomorphic in plumage‐coloration. In this species, sexual and territorial call behaviour is mainly performed at dawn and dusk, during which a white patch on the throat is repeatedly exposed at each call. We measured the total light reflectance of the feathers of this badge in 39 eagle owl specimens from museum collections. We found seasonal variability and sexual dichromatism in the brightness of the plumage badge. The total reflectance of this trait peaked during the territorial‐mating period. Moreover, females showed higher values of brightness than males, in agreement with the reversed body size dimorphism present in this and many other raptor species. Finally, female but not male body size was positively correlated with white badge reflectance.     

Males in seemingly female‐like plumage do not mimic females: UV reflectance reveals temporal cryptic dimorphism in a manakin species exhibiting delayed plumage maturation

Manakins (Pipridae) are neotropical birds that usually exhibit delayed plumage maturation (DPM). Thus, while plumage of most adult male manakins is brightly conspicuous, subadult males and females are basically dull‐olive green. Although sexual dichromatism in some bird species may be evident only through UV reflectance, this phenomenon, known as hidden sexual dichromatism, has not been previously studied in manakins to compare subadult males and females. Within this framework, we carried out spectrophotometric analyses in searching for hidden sexual dichromatism in the white‐bearded manakin Manacus manacus, through comparison of UV spectra in females and subadult males in green plumage. Our results revealed UV reflectance in both sexes in green plumage. Moreover, we found UV spectral differences in homologous color patches between sexes, particularly at belly. Since the observed differences may allow intraspecific sex recognition of individuals in green plumage, our results do not support the female‐mimicry hypothesis to explain delayed plumage maturation in the white‐bearded manakin. Although our findings dismiss the female mimicry hypothesis, we cannot state whether these results support the non‐mutually exclusive cryptic and status signaling hypotheses. We propose then, that dull coloration of subadult males may serve both as a cryptic trait and to limit the energetic costs of acquiring the adult plumage before sexual maturity. Meanwhile, differential UV color traits between sexes in green plumage may allow adult males to avoid unnecessary energy expenditures in courtship displays in the presence of males near leks, and to selectively focus their the courtship displays on females. In accordance with the status signaling hypothesis, subadult males can be recognized both as males and subordinates and consequently may practice courtship displays without suffering aggressions by adult males. Our results highlight the importance to include a wider range of spectrophotometric information analyses for testing hypotheses regarding delayed plumage maturation.     

Alula size signals male condition and predicts reproductive performance in an Arctic‐breeding passerine

While studies of achromatic plumage signaling are scarce relative to chromatic ornaments, achromatic ornaments have the potential to act as an efficient form of visual communication due to the highly conspicuous contrast between black and white body regions. Recently, achromatic plumage reflectance has been shown to indicate condition, yet the condition‐dependence of achromatic patch size remains unstudied. Here we show the first evidence that alula size, an achromatic plumage patch, has the potential to signal a male’s condition and predict reproductive performance. In Arctic‐breeding snow buntings Plectrophenax nivalis, the size of the alula simultaneously predicted pre‐breeding physiological health and the number of offspring produced, through an intermediate variable (lay date). Snow buntings appear to pair assortatively; males and females arriving earlier pair together, and changes in body condition over the breeding season are positively related within pairs. We suggest that simple achromatic plumage patches, like alula size, have the potential to act as condition‐dependent signals. Consequently, females may benefit from assessing these signals to reliably evaluate a male’s condition and reproductive potential as a means of maximizing their reproductive success.     

Strategic concealment of sexual identity in an estrilid finch

One explanation for the evolution of sexual monomorphism is the sexual indistinguishability hypothesis, which argues that in group-living species individuals might benefit by concealing their sex to reduce sexual competition. We tested this hypothesis in long-tailed finches Poephila acuticauda. Males and females could not be reliably distinguished morphologically or by analysis of the reflectance spectra (300 to 700 nm) from the plumage and bill. Males seemed unable to distinguish the sex of an unfamiliar individual in the absence of behavioural cues; they were equally likely to court and copulate with unfamiliar males and females but rarely courted familiar males. Here we report the first experimental evidence that sexual monomorphism enables strategic concealment of sex. Males were more likely to reveal their sex when faced with a solitary unfamiliar individual than a group of unfamiliar individuals. When encountering an unfamiliar male that revealed his sex, subordinate males were more likely to conceal their sex than dominant males.     

Ultraviolet reflection and female mate choice in the pied flycatcher, Ficedula hypoleuca

In pied flycatchers females seem to prefer male territory quality rather than male characteristics, and the results of female mate choice experiments are divergent. In this outdoor aviary study, we examined how altering the ultraviolet reflection of males affects female mate choice behaviour. We chose pairs of males with similar human-visible dorsal colour and morphological traits. We then reduced the proportional ultraviolet reflectance in one male with sunscreen chemicals. The other male was treated with a chemical that slightly increased the ultraviolet reflectance of the plumage. In the experiment females clearly preferred males with slightly increased ultraviolet reflection. Our results indicate that pied flycatcher females use ultraviolet cues for mate choice when the effect of territory quality is controlled for. The results give us new information about a possible mechanism of mate assessment in this species, and indicate the importance of colour cues in avian mate choice behaviour.     

The role of female dominance hierarchies in the mating behaviour of mosquitofish

While studies of sexual selection focus primarily on female choice and male-male competition, males should also exert mate choice in order to maximize their reproductive success. We examined male mate choice in mosquitofish, Gambusia holbrooki, with respect to female size and female dominance. We found that the number of mating attempts made by a male was predicted by the dominance rank of females in a group, with dominant females attracting more mating attempts than subordinates. The number of mating attempts made by males was independent of the female size. The observed bias in the number of mating attempts towards dominant females may be driven either by straightforward male mate choice, since dominance and female fecundity are often closely related, or via the dominant females mediating male mating behaviour by restricting their access to subordinate females.     

Sexual Selection in Black Morph Girardinus metallicus (Pisces: Poeciliidae): Females Can Spot a Winner (But We Cannot)

Male–male competition may interfere with the ability of females to choose mates by interrupting courtship or by favoring highly aggressive males who may damage females during mating attempts. Alternatively, females may benefit by mating with dominant males, and female choice and male–male competition may therefore act in unison. The same traits, including aggressiveness, may indicate male quality to females and to rivals. We investigated sexual selection in the black morph of the endemic Cuban poeciliid fish, Girardinus metallicus, to ascertain the links between morphological and behavioral traits and success in intra‐ and intersexual selection. Males conspicuously exhibit their black ventral surface and gonopodium to females during courtship. Dichotomous choice tests revealed female association preferences for certain males, and those same males were more successful in monopolizing access to females when the fish were allowed to directly interact. Dominant males followed, courted, and copulated with females more than subordinate males within a pair, and it appears that females could either assess dominance based on cues we did not measure, or could influence subsequent mating success by their behavior during the dichotomous choice trials. There was an interaction between black status (i.e., whether the male in each pair had more or less ventral black coloration than the other male in that pair) and dominance, such that low‐black dominant males courted early and then shifted to following females, whereas high‐black dominant males courted far more later in the observation period. These results hint at the importance for sexual selection of the interplay between a static morphological trait (black coloration) and a dynamic behavioral trait (aggressiveness), but the functional significance of the courtship display remains a mystery.     

The influence of a male's dominance status on female choice in Syrian hamsters

This study evaluated the importance of dominance status on mate selection in Syrian hamsters. In one experiment, sexually receptive females were allowed to choose between tethered males which differed in status. The choice was consistently in favor of the dominant male. The female spent more time in lordosis in the presence of the dominant male. The dominant also obtained a significantly greater number of intromissions. A second experiment investigated whether prior familiarization was essential to permit a female to express mating preferences in a situation where she was free to interact with three males. Again, the dominants were most often preferred and obtained greater sexual access to the female. Prior familiarization or extensive contact with the males was not necessary to support the female's selection of the dominant as a mating partner. The choice occurred quickly, generally within 5 min after contacting the males. Although females did mate with the subordinates, this typically occurred late in the tests. The significance of these data with respect to mate choice and probable paternity effects are discussed.     

Brightness variability in the white badge of the eagle owl Bubo bubo

The application of modern spectrometry to the study of avian colour variability has revealed ignored patterns of colour variation such as male-biased sexual dichromatism and seasonal variability in the plumage. However, the variation in the achromatic property of such traits, that is in the total light reflectance of the spectrum (i.e., brightness), has commonly been overlooked. The evolution of signals based on brightness should be favoured in those species that are active when light is scarce, i.e. at dawn and dusk. The eagle owl Bubo bubo is monogamous and apparently monomorphic in plumage-coloration. In this species, sexual and territorial call behaviour is mainly performed at dawn and dusk, during which a white patch on the throat is repeatedly exposed at each call. We measured the total light reflectance of the feathers of this badge in 39 eagle owl specimens from museum collections. We found seasonal variability and sexual dichromatism in the brightness of the plumage badge. The total reflectance of this trait peaked during the territorial-mating period. Moreover, females showed higher values of brightness than males, in agreement with the reversed body size dimorphism present in this and many other raptor species. Finally, female but not male body size was positively correlated with white badge reflectance.     

Female dominance rank and behavior during artificial menstrual cycles in social groups of rhesus monkeys (Macaca mulatta)

We recently reported (Michael & Zumpe: American Journal of Primatology 15:157–170, 1988) evidence for the hypothesis that copulations by pairs of rhesus monkeys are linked more closely to the ovulatory phase of the female's menstrual cycle when a male has access to several females in different cycle phases rather than access to females in the same cycle phase. We have now used data from this earlier study to assess the role of female dominance. Artificial menstrual cycles were induced with hormone injections in two of four ovariectomized females in eight social groups (each consisting of one male and four females). The artificial cycles were either synchronized or offset by 7-day increments. There were differences in the behavioral interactions of the higher-ranking (dominant) and lower-ranking (subordinate) hormone-treated females in each group. The amount of male sexual activity received by dominant and subordinate females differed little, but dominant females initiated significantly more sexual activity (P<.0001) than did subordinate females. Dominant females were significantly more aggressive (P<.0001) toward other females and received more direct and redirected aggression from males. Males generally initiated and maintained more social contacts with dominant than with subordinate females, and dominant females were less active than subordinates in initiating and maintaining social interactions with males. The findings provide new evidence for both female mate competition and mate choice by males for higher-ranking females.     

Male competition and female choice interact to determine mating success in the bluefin killifish

Whether male competition and female choice act in concert, independently,or in opposition is a critical issue for understanding sexualselection. In complex social systems, the outcomes of pairwiseinteractions may not be accurate indicators of how sexual selectionemerges. We investigated how female choice and male competitioninteract in the bluefin killifish, Lucania goodei, in a 3-stagedexperiment where 1) females could choose between 2 males, 2)those males could interact in the presence of that female, and3) females and males could freely interact and spawn. In thepairwise stages (1 and 2), females displayed pronounced preferencesbetween males and male competition produced a distinctly dominantindividual. None of the morphological traits, including color,measured in males were associated with either female preferenceor male dominance. When all 3 fish interacted (stage 3), maleactivity level was the sole predictor of spawning success. Maleswith elevated activity levels were more aggressive toward malesand females, exhibited intensified courtship, and obtained morespawns. Female preference did not predict the number of spawnswith a male, but it did predict her latency to spawn; femalesspawned more quickly with preferred males. Thus, male competitionand female choice interact to determine reproductive success,but there is evidence for conflict and a cost to females ofassociating with dominant males. Reproductive success in thisspecies is not easily predicted from simple measures of morphologyor female preference and is influenced by complex social interactions,both between males, and between males and females.     

Do Female Mandrills Prefer Brightly Colored Males?

Although secondary sexual adornments are widespread in male primates, few studies have examined female choice for these characters. Mandrills (Mandrillus sphinx) present an extreme example of sexual dimorphism, with males exhibiting an array of striking adornments. The most dominant adult male in a group exhibits the brightest and most extensive red coloration, while the other males are less brightly colored. I examined whether female mandrills prefer brightly colored males using data on periovulatory sexual behavior during the 1996 mating season for all males 8 years old (n = 5) and all parous females (n = 9) in a semifree-ranging colony at CIRMF, Gabon. Brightness of male coloration is significantly positively correlated with time spent within 2 m of females, female responsibility for proximity, number of sexual presentations received, % approaches accepted by females, and % inspections with which females cooperated. Females also groomed only the brightest male. Behaviors indicating female preference are not correlated significantly with male dominance rank, and partial correlations confirm that the influence of male color on female behavior is stronger than that of male rank. With the influence of male dominance rank controlled, correlation coefficients between female behaviors and male mating success are high and positive. In further support of the hypothesis that females show mate choice for brightly colored males, independent of dominance rank, I report an unusual case wherein the alpha male fell in rank without loss of coloration. He experienced no significant change in female responsibility for proximity, sexual presentations received, or female reaction to approaches or inspections, though he was no longer observed to mate. Accordingly, female mandrills attend to differences in male secondary sexual characters and favor brightly colored males. As brightly colored males are also dominant this reinforces the influence of male-male competition on male reproductive success and may explain the very high reproductive skew in mandrill males and their extraordinary appearance.     

Plumage Colouration,Age, Testosterone and Dominance in Male Red Bishops (Euplectes orix): A Laboratory Experiment

Visual signals such as plumage characteristics in birds often play a key role in the establishment of dominance in contests by acting as a badge of status that can be used to assess individual fighting ability. We studied the role of plumage colouration in males of the red bishop (Euplectes orix), a sexually dimorphic and polygynous weaverbird species occurring in sub‐Saharan Africa and breeding in dense colonies around water. Males are highly territorial and often engage in competition over limited resources such as breeding sites and potential mates. By experimentally staging male–male contests over a limited food source, we wanted to determine whether the orange–red breeding plumage in this species serves as a dominance signal between individuals, with males with redder plumage being dominant over those with duller plumage. In the first set of experiments, we staged contests between unfamiliar and unmanipulated males. The setup of the second set of experiments was identical to the first, with the exception that those males with the lowest chroma and hue values had their plumage experimentally reddened within the range of the natural variation. In addition to plumage colouration, we incorporated testosterone levels, body condition and age into the analysis of factors contributing to contest outcome. Our results show a consistent and strong age effect in both sets of experiments, which seems to be independent of plumage colouration, testosterone and body condition. This suggests that in the red bishop, the outcome of male–male competitions over limited resources is determined by age‐related acquired experience rather than by status signalling through plumage colouration.     

The evolution of postpairing male mate choice

An increasing number of empirical studies in animals have demonstrated male mate choice. However, little is known about the evolution of postpairing male choice, specifically which occurs by differential allocation of male parental care in response to female signals. We use a population genetic model to examine whether such postpairing male mate choice can evolve when males face a trade‐off between parental care and extra‐pair copulations (EPCs). Specifically, we assume that males allocate more effort to providing parental care when mated to preferred (signaling) females, but they are then unable to allocate additional effort to seek EPCs. We find that both male preference and female signaling can evolve in this situation, under certain conditions. First, this evolution requires a relatively large difference in parental investment between males mated to preferred versus nonpreferred females. Second, whether male choice and female signaling alleles become fixed in a population versus cycle in their frequencies depends on the additional fecundity benefits from EPCs that are gained by choosy males. Third, less costly female signals enable both signaling and choice alleles to evolve under more relaxed conditions. Our results also provide a new insight into the evolution of sexual conflict over parental care.     

Familiarity adds to attractiveness in matters of siskin mate choice

There is currently considerable controversy in evolutionary ecology revolving around whether social familiarity brings attraction when a female chooses a mate. The topic of familiarity is significant because by avoiding or preferring familiar individuals as mates, the potential for local adaptation may be reduced or favoured. The topic becomes even more interesting if we simultaneously analyse preferences for familiarity and sexual ornaments, because when familiarity influences female mating preferences, this could very significantly affect the strength of sexual selection on male ornamentation. Here, we have used mate-choice experiments in siskins Carduelis spinus to analyse how familiarity and patterns of ornamentation (i.e. the size of wing patches) interact to influence mating success. Our results show that females clearly prefer familiar individuals when choosing between familiar and unfamiliar males with similar-sized wing patches. Furthermore, when females were given the choice between a highly ornamented unfamiliar male and a less ornamented familiar male, half of the females still preferred the socially familiar birds as mates. Our finding suggests that male familiarity may be as important as sexual ornaments in affecting female behaviour in mate choice. Given that the potential for local adaptation may be favoured by preferring familiar individuals as mates, social familiarity as a mate-choice criterion may become a potential area of fruitful research on sympatric speciation processes.     

RESEARCH PAPER: Multiple Sexual Ornamentation Signals Male Quality and Predicts Female Preference in Minnows

Sexual ornamentation often consists of multiple components. Different sexual signals may indicate different aspects of mate quality or reflect quality in different time scales. On the other hand, same signals can have a dual function and are used both in male–male competition and courtship. Many fish species are capable of rapidly altering their colouration (ephemeral colour changes), but this capability is usually ignored in sexual selection studies. Here, we used experimentally manipulated social environments to study the ephemeral colour changes in multicomponent sexual signals of male minnows (Phoxinus phoxinus) during male–male competition and female choice. We found that the dominant males courted the females more actively and had redder and/or darker skin colouration than the subordinate males. Furthermore, darkness difference between subordinate and dominant males increased in the presence of female, which suggests that the male–male competition may increase the honesty of signalling and thus facilitate female choice. In support of this hypothesis, females had a strong behavioural preference towards the more colourful males, which may indicate female choice. As colourful males often had a higher social status than paler individuals, it is possible that females base their preference on male status, not only the colouration per se. In any case, our results suggest that sexual ornamentation of male minnows may signal status, courting activity and superior quality of the males and that these signals may have a dual function in both male–male competition and female choice. Females preferred different ornamental traits (dark and red colour patterns) relatively equally, indicating that mate choice is based on multiple cues.     

Structural plumage colour and parasites in satin bowerbirds Ptilonorhynchus violaceus: implications for sexual selection

We investigated whether variation in structural plumage coloration in satin bowerbirds, Ptilonorhynchus violaceus , could reveal the intensity of infection from parasites, as predicted from models of parasite-mediated sexual selection (PMSS). To do this, we captured adult male, female, and juvenile male satin bowerbirds in Queensland, Australia, and objectively measured individual plumage reflectance from four body regions using a spectrometer. We quantified both ectoparasite load and the intensity of infection from blood parasites. In iridescent blue adult males, plumage reflectance is unimodal, with a single peak in the ultraviolet, while in greenish females and juveniles, plumage reflectance is bimodal, with peaks in both the ultraviolet and green portions of the spectrum. In adult males, the intensity of infection from blood parasites was best predicted by plumage brightness (total reflectance), with brighter males having fewer parasites. Similarly, juvenile males exhibiting greater UV chroma (proportion of reflectance in the UV) had fewer blood parasites. Our findings support a key prediction of PMSS models and provide the first evidence that a structural colour ornament can signal the intensity of infection from blood parasites.