Aggressive Behavior and Courtship Rejection in Brightly and Plainly Colored Female Keeled Earless Lizards (Holbrookia propinqua)

Relationships between bright secondary sexual coloration and behavior were studied in female Holbrookia propinqua, which develop striking orange and yellow colors during the breeding season. In tethered introduction studies, brightly colored females performed aggressive courtship rejection behaviors toward conspecific males; plainly colored females were not aggressive toward males, but attempted to avoid them. Responses of females of the two color patterns toward conspecific females of both color phases were not detectably different. Experimental introductions of lizards with coloration modified by paint showed that females of both color patterns recognize any other lizard bearing the bright female colors as female, regardless of actual sex. Both the orange and yellow components were shown to contribute to sex recognition. The yellow component alone allowed accurate sex identification, but only half the females responded to males painted with only the orange female component as if they were females. Because females did not behave differently toward other females on the basis of coloration, the hypothesis that bright coloration evolved as an adaptive signal between females is rejected. The dark ventrolateral stripes of male and plainly colored females did not appear to affect intraspecific social responses by females.     

Carotenoid-based ornaments of female and male American goldfinches (Spinus tristis) show sex-specific correlations with immune function and metabolic rate

Conspicuous ornamentation has been linked to immunological and physiological condition in males of many species. In species where both sexes are ornamented, it is unclear whether the signal content of ornaments differs between males and females. We examined the immunological and physiological correlates of carotenoid-based bill and plumage ornamentation in American goldfinches Spinus tristis, a species in which bright orange bills are sexually monomorphic but yellow plumage is sexually dimorphic during the breeding season. Because bill color is dynamic over short periods while plumage color is static over longer time frames, we tested whether these signals have the potential to provide temporal information about immunity and condition. In both sexes, bill color (but not plumage color) was negatively related to leukocyte differential, a measure of recent stress, while plumage color (but not bill color) was positively related to resting metabolic rate. In females, bill color also positively correlated with immunoglobulin Y, a component of acquired immunity, while plumage color positively predicted natural antibody levels, a component of innate immunity. In males, neither bill color nor plumage color predicted immune function, suggesting that the mechanisms underlying these signals vary with sex. Our results demonstrate that dynamic signals such as bill coloration do not merely reflect the same information provided by static signals but that these two classes of signal provide information about different temporal aspects of phenotypic quality. Furthermore, our results indicate that a signal expressed in both sexes has the potential to provide different information depending on the sex of the bearer.     

Status signalling in male but not in female Eurasian Tree Sparrows Passer montanus

Male ornaments, such as plumage coloration, frequently serve as signals. The signalling function of similar ornaments in females has, however, received much less attention despite the fact that conspicuousness of their ornaments is often comparable to those of males. In this study we tested the signalling function of a plumage trait present in both sexes in the Eurasian Tree Sparrow Passer montanus. The black throat patch has been repeatedly found to have a signal function in the closely related House Sparrow Passer domesticus, where only males bear the ornamental trait. However, the function of the black throat patch in the females of Passer species that have sexually monomorphic ornament expression has never, to our knowledge, been considered. We investigated the outcomes of aggressive encounters in foraging flocks of free‐living Tree Sparrows, and assessed whether throat patch size and measurements of body size predicted fighting success in these flocks. We found that male throat patch size predicted fighting success against both male and female opponents. However, female throat patch size did not correlate with fighting success against either sex. Among the morphological traits studied, wing length was the best predictor of fighting success in females. Our findings suggest a status signalling function of throat patch size in males but not in females, although further experimental studies are necessary to corroborate these correlative results.     

Wake up and smell the conflict: odour signals in female competition

Odour signals used in competitive and aggressive interactions between males are well studied in the context of sexual selection. By contrast, relatively little is known about comparable signals used by females, despite current interest in the evolution of female ornaments and weaponry. Available evidence suggests that odour signals are important in competitive interactions between female mammals, with reductions or reversals of male-biased sexual dimorphism in signalling where female competition is intense. Scent marking is often associated with conflict between females over access to resources or reproductive opportunities. Female scent marks may therefore provide reliable signals of competitive ability that could be used both by competitors and potential mates. Consistent with this hypothesis, we report that aggressive behaviour of female house mice is correlated with the amount of major urinary protein (MUP) excreted in their urine, a polymorphic set of proteins that are used in scent mark signalling. Under semi-natural conditions, females with high MUP output are more likely to produce offspring sired by males that have high reproductive success, and less likely to produce offspring by multiple different sires, suggesting that females with strong MUP signals are monopolized by males of particularly high quality. We conclude that odour signals are worthy of more detailed investigation as mediators of female competition.     

Dominance, Status Signals and Coloration in Male Mandrills (Mandrillus sphinx)

Where individuals contest access to a resource, escalated physical fighting presents a risk to all involved. The requirement for mechanisms of conflict management has led to the evolution of a variety of decision rules and signals that act to reduce the frequency of aggression during competitive encounters. We examined strategies of conflict management in male mandrills (Mandrillus sphinx) living in two semi‐free‐ranging groups in Gabon. Adult male mandrills are large (31 kg), with long canines, making the costs of conflict potentially very high. We found that males formed dominance hierarchies, but that male–male relationships were characterized by avoidance, appeasement and ignoring. Fights were rare, but could result in death. Examination of the relationship between dominance and signaling showed that males use facial and gestural signals to communicate dominance and subordinance, avoiding escalated conflict. Male mandrills also possess rank‐dependent red coloration on the face, rump and genitalia, and we examined the hypothesis that this coloration acts as a ‘badge of status’, communicating male fighting ability to other males. If this is the case, then similarity in color should lead to higher dyadic rates of aggression, while males that differ markedly should resolve encounters quickly, with the paler individual retreating. Indeed, appeasement (the ‘grin’ display), threats, fights and tense ‘stand‐off’ encounters were significantly more frequent between similarly colored males, while clear submission was more frequent where color differences were large. We conclude that male mandrills employ both formal behavioral indicators of dominance and of subordination, and may also use relative brightness of red coloration to facilitate the assessment of individual differences in fighting ability, thereby regulating the degree of costly, escalated conflict between well‐armed males.     

The relationship between plumage coloration and aggression in female tree swallows

Intrasexual competition is an important selective force that can favor the evolution of honest signals of fighting ability. Research has focused predominantly on male birds, but many female birds also possess plumage ornaments that could mediate the outcome of competitive interactions. We examined the relationship between blue and white structural coloration and aggression in female tree swallows Tachycineta bicolor. Tree swallows are secondary cavity nesters and females show delayed plumage maturation which may be related to intense competition for nest sites. We compared plumage reflectance of second‐year (SY) and after‐second year (ASY) females, and within‐individual changes in plumage reflectance of ASY females in two successive years. We assessed aggression by placing a caged SY female 2 m from the nest box of an ASY female and quantified the ASY female's response in relation to her own plumage coloration. We found substantial differences in plumage reflectance of SY and ASY females, but found that ASY females became greener with duller white coloration in the second year. This may be due to poor weather that made reproduction particularly costly in the first year of the study and suggests coloration is influenced by condition. We found no relationship between dorsal coloration and aggressiveness. Rather, brighter white females spent more time on their nest box and within 2 m of the intruder than females with dull breasts. Our findings suggest that brighter white ASY females may perceive a SY female as less of a threat or that there may be a trade‐off associated with aggression and parental care that is related to white brightness.     

Weaponry, color, and contest success in the jumping spider Lyssomanes viridis

Weaponry and color badges are commonly theorized to function as visual signals of aggressiveness or fighting ability. However, few studies have supported a signaling function of weaponry, and the role of color in invertebrate competitive interactions remains virtually unexplored. Jumping spiders (Salticidae) make excellent invertebrate models for studying weaponry and color because males of many species are colorful and possess exaggerated chelicerae, which are used as weapons in escalated contests. To determine whether color or weaponry might function as visual signals in male-male competitions, we investigated relationships between contest success, cheliceral length, and red coloration in Lyssomanes viridis. Males having longer chelicerae than their opponents were significantly more likely to win (p=0.0008). Males who won, despite being smaller than their opponents, had significantly less red chelicerae than their opponents (p=0.01). Male and female cheliceral length, as well as foreleg length, correlated tightly with body size. Cheliceral and foreleg length showed significantly stronger positive allometry in males than in females. We conclude that male chelicerae and forelegs are under strong positive selection for their use in physical fights and/or as visual signals of fighting ability.     

Genes, copying, and female mate choice: shifting thresholds

Recent experimental work on guppies (Poecilia reticulata) hasexamined the strength of genetic and cultural (copying) factorsin determining female mate choice. Using females from a populationwith a heritable preference for the amount of orange body colorpossessed by males, prior work discovered that a threshold differencein orange color among males existed below which females wouldchoose a less orange male if they observed another female choosethat male, but above which they consistently preferred the moreorange of the males, regardless of whether they viewed anotherfemale prefer the less orange male. I tested whether this thresholdcan be shifted by increasing the amount of mate-copying informationavailable to a female. I demonstrate that when a female hasthe opportunity to see two different model females independentlyprefer the less orange of two males or a single female neara drab male for a longer period of time (twice as long as inprior work), the observer female prefers this drab male evenwhen males dramatically differ in orange coloration.     

Preferences of female American goldfinches (Carduelis tristis) for natural and artificial male traits

We conducted two experiments to determine mating preferencesof female American goldfinches (Carduelis tristis). In the firstexperiment, females were allowed to choose among four naturallyvarying males. In the second, females chose among four maleswearing either orange, blue, yellow, or no color bands. In experiment1, females performed more courtship with males that had significantlybrighter bills and were brighter overall. In experiment 2, femalespreferred males wearing orange bands over unhanded males. Blue-bandedmales ranked second and yellow third, but neither was preferredsignificantly more often than the unbanded males. A female'spreference for a particular band color was weakly associatedwith her own bill and plumage measures; females with brighterthroat feathers and yellower bills preferred orange bands toblue bands. The results suggest that female preferences fororange and yellow may be functional and that females may alsohave aesthetic preferences.     

Evidence for sexual selection on structural plumage coloration in female eastern bluebirds (Sialia sialis)

Although the function of ornamental traits in males has been the focus of intensive research for decades, expression of such traits in females has received much less study. Eastern bluebirds (Sialia sialis) display structurally based ultraviolet/blue and melanin-based chestnut plumage, and in males this plumage coloration is related to both reproductive success and competitive ability. Compared to males, female bluebirds show a subdued expression of blue and chestnut ornamental coloration, and we used a combination of an aviary nutritional-stress experiment and four years of field data to test the hypothesis that coloration functions as a signal of female quality. First, we tested the effect of food intake on expression of structural and melanin coloration in female eastern bluebirds to determine whether structural or melanin coloration are condition-dependent traits. Females that were given ad libitum access to food displayed more ornamented structural coloration than females on a food-restricted diet, but there was no effect of the experiment on melanin ornamentation. Second, we used field data to assess whether female ornamentation correlated with measures of mate quality and parental effort. The structural coloration of females predicted first egg date, maternal provisioning rates, and measures of reproductive success. These data indicate that structural coloration is dependent on nutritional condition and suggest that sexual selection is acting on structurally based plumage coloration in female eastern bluebirds.     

Female Preference for Sympatric vs. Allopatric Male Throat Color Morphs in the Mesquite Lizard (Sceloporus grammicus) Species Complex

Color polymorphic sexual signals are often associated with alternative reproductive behaviors within populations, and the number, frequency, or type of morphs present often vary among populations. When these differences lead to assortative mating by population, the study of such polymorphic taxa may shed light on speciation mechanisms. We studied two populations of a lizard with polymorphic throat color, an important sexual signal. Males in one population exhibit orange, yellow, or blue throats; whereas males in the other exhibit orange, yellow, or white throats. We assessed female behavior when choosing between allopatric and sympatric males. We asked whether females discriminated more when the allopatric male was of an unfamiliar morph than when the allopatric male was similar in coloration to the sympatric male. We found that female rejection of allopatric males relative to sympatric males was more pronounced when males in a pair were more different in throat color. Our findings may help illuminate how behavioral responses to color morph differences between populations with polymorphic sexual signals contribute to reproductive isolation.     

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.     

Female preference for dominant males in the Mexican mojarra cichlid fish,Cichlasoma istlanum

It is widely assumed that female preference and male competition operate simultaneously during sexual selection. Dominance is likely an honest indicator of male quality, and females can identify and choose the dominant male to reproduce with individuals with greater competitive abilities, thus improving the quality and competitiveness of their offspring. In this context, few studies have investigated female preference in relation to male fighting ability. The Mexican mojarra, Cichlasoma istlanum, is a cichlid species native to the Balsas River basin. It is territorial during reproduction and provides parental care. Males commonly engage in territorial defence, whereas females care directly for offspring. This study examined whether females prefer dominant males that exhibit more aggressive behaviour. The authors conducted experiments using groups of two males and one female to test competitive ability in males and female preference. They also quantified the time during which the female associated with the dominant male and the subordinate male after observing the outcome of a fight between the two males. They found that Mexican mojarra females preferred dominant males and that the time females spent associating with males was positively related with their aggressive behaviours during competition. These results indicate that dominant males were more attractive than subordinate males to female Mexican mojarra. The relationship between female preference and male dominance in the Mexican mojarra demonstrates the importance of male competitive ability for future parental care in reproduction.     

Signal content of red facial coloration in female mandrills (Mandrillus sphinx)

Studies of secondary sexual ornamentation and its maintenance by sexual selection tend to focus on males; however, females may also possess showy ornaments. For example, female mandrills possess facial coloration that ranges from black to bright pink. We used fortnightly photographs of 52 semi-free-ranging females aged above 3years over 19 months to evaluate whether colour conveys information concerning female competitive ability, reproductive quality, age or reproductive status. Colour was not related to female rank or quality (body mass index, age at first birth or mean inter-birth interval); however, colour did increase significantly with age and primiparous females were darker than multiparous females. Colour may therefore signal reproductive quality, as younger females are less fertile and produce smaller offspring. Colour was brighter during the follicular phase than during the luteal phase, suggesting that it may signal fertility. Colour also varied across gestation and peaked at four and eight weeks post-parturition, suggesting that it may signal approaching parturition and lactation. Future studies should examine the relationship between colour and the menstrual cycle in more detail, the hormonal basis of female colour, and determine experimentally whether mandrills of both sexes attend to differences in colour between and within females.     

Bill Color Preferences of Zebra Finches

Bill color varies with age and sex in zebra finches. Among birds of similar age and condition, males' bills tend to be redder and darker than those of females, but there is overlap in the phenotypic expression of the sexes. The bills of young birds are paler and less red than those of older birds. There is also interindividual variation within age and sex class. Experiments were performed to measure heterosexual and isosexual (= same sex) preferences of finches. Females preferred to associate with males with the reddest, brightest bills; they even preferred males whose bills were exaggerated through color applications of non-toxic marking pen. Males preferred to associate with females with bill colors in the middle of the phenotypic range. Females thus have “directional” preferences for male bill color, whereas male preference is “stabilizing” with regard to female bill color. In isosexual tests, neither sex showed a consistent preference for particular bill colors. Both sexes, however, displayed a tendency toward individual variability in preference. Bill color appears to be more important in heterosexual than in isosexual interactions. Several authors have recently suggested that organisms prefer brightly colored mates because bright coloration indicates superior physical condition. Results reported here do not support this hypothesis. Alternative functional explanations for the observed preferences are that bill color signals mating status, age or reproductive value. None of these appears to be a cogent explanation for the trends. Preferences do not appear to result from sexual imprinting. The possibility that the preferences are aesthetic and non-functional is discussed.     

Sexual Signals of the Male American Goldfinch

Male American goldfinches (Carduelis tristis) exhibit conspicuous yellow plumage, orange bills, and black caps during the breeding season. These secondary sexual characteristics may serve as criteria by which females evaluate males as potential mates because the traits vary among individuals and are likely to be costly. We quantified plumage and bill color and cap characteristics of wild, free‐ranging American goldfinches during the breeding season and tested for relationships between those features, body condition and individual genetic diversity in males. Overall, male and female goldfinches were highly sexually dichromatic, with plumage saturation and brightness and bill brightness contributing strongly to the dimorphism. Body condition decreased significantly with Julian date, even over the 2‐wk period immediately prior to the onset of nesting during which we collected our color and cap measurements. Principal components describing color of the back and the bill significantly predicted date‐corrected body condition based on quadratic regressions, suggesting that there is reliable information in back and bill color that females could use when choosing mates. In a subset of captive males, we found that bill hue faded from orange to yellow within 24 h of capture, suggesting that bill color may reflect short‐term changes in health status or carotenoid availability. Individual genetic diversity based on a panel of eight microsatellite loci was correlated with back brightness and perhaps with cap symmetry. Based on the results of this field study, ornamentation of male American goldfinches appears to signal both long‐ and short‐term aspects of phenotypic quality.     

Social costs and development of nuptial coloration in male Psammodromus algirus lizards: an experiment

In the lizard Psammodromus algirus, larger and older males show orangenuptial coloration on most of the head and are dominant oversmaller and younger, albeit sexually mature, males which donot show such extensive nuptial coloration. This raises thequestion of why young, small males delay the development ofnuptial coloration until a later breeding season. We tested thehypothesis of social costs by manipulating the color of thehead of small males. The results of agonistic interactions suggestedthat small males may pay a cost in terms of being punished bylarge males. Small males with heads painted orange were stillrecognized as small by other small males, suggesting that theywould not gain in social status relative to normal, dull, small males.We also manipulated the coloration of large males. Small malesshowed a similar response toward all large males, independentof coloration. This suggests that in short-distance communication,males used other cues, such as body size and behavior, whenjudging fighting ability. In staged experiments without malecompetition, female acceptance of matings was influenced bymale body size but not by coloration because large males weremore successful in obtaining matings than were small males,and within each age/size category there was no difference inmating success between experimental and control males.     

Reproductive correlates of plumage coloration of female Mountain Bluebirds

Many studies have shown that the plumage coloration of male birds can act as an honest signal of quality, indicating benefits that a female could gain from pairing with a specific male. In some species, females also display ornamental plumage, but less is known about the function and potential adaptive significance of female coloration because most research has focused on male coloration. Male Mountain Bluebirds (Sialia currucoides) display full body, ultraviolet (UV)‐blue plumage, whereas female plumage is more subdued, with blue color focused on the rump, wing, and tail. During the 2011 and 2012 breeding seasons (May–July) near Kamloops, BC, Canada, we examined coloration of the rump and tail of female Mountain Bluebirds to determine if their plumage could act as an indicator of direct reproductive benefits (e.g., enhanced parental care or reproductive success) to potential mates. We found no relationship between female plumage coloration and either provisioning rate or fledging success. However, female coloration varied with age, with after‐second‐year (ASY) females having brighter, more UV‐blue tail feathers than second‐year (SY) females. In addition, ASY females with brighter, more UV‐blue tails had larger clutches. We also observed positive assortative mating by tarsus length. Because previous work with other species suggests that female body size may be a good predictor of breeding success, males could potentially benefit from pairing with larger females. However, reproductive success did not vary with female size in our study. Although our evidence that structural plumage coloration of female Mountain Bluebirds is a signal of direct reproductive benefits for males (e.g., higher reproductive success) is limited, our results (i.e., ASY females with brighter tails than SY females, and ASY females with brighter tails having larger clutches) do suggest the potential for sexual selection to act on female coloration.     

Eggshell coloration and its importance in postmating sexual selection

Avian eggshell color seems to fulfill multiple functions, some of them being structural and others signaling. In this study, we tested whether or not eggshell coloration may play a role in sexual selection of Tree Sparrows (Passer montanus). According to the “Sexually selected eggshell coloration” hypothesis, eggshell coloration signals female, egg or chick quality and males adjust parental investment according to this signal. Eggs of this species are covered with brown spots and patches, and variation between clutches is high. We found that eggshell coloration correlates with both protoporphyrin and biliverdin, but protoporphyrin concentrations are ten times higher. Eggshell coloration reflects egg and offspring quality, but not female quality. Thus, eggshell coloration may signal female postmating investment in offspring rather than female quality. Furthermore, differential allocation in terms of maternal investment is supported by the fact that females lay more pigmented clutches when mated to males with bigger melanin‐based ornaments relative to their own. Moreover, males invested proportionally more to chicks that hatched from more pigmented clutches. Our correlative results thus seem to support a role of sexual selection in the evolution of eggshell coloration in birds laying brown eggs, pigmented mainly by protoporphyrin.     

Ornamental bill color rapidly signals changing condition

Ornamental bill color is postulated to function as a condition‐dependent signal of individual quality in a variety of taxonomically distant bird families. Most red, orange, and yellow bill colors are derived from carotenoid pigments, and carotenoid deposition in ornamentation may trade off with their use as immunostimulants and antioxidants or with other physiological functions. Several studies have found that bill color changes in response to physiological perturbations, but how quickly such changes can occur remains unclear. We tested the hypothesis that carotenoid‐based orange bill color of American goldfinches Spinus tristis responds dynamically to rapid changes in physiological stress and reflects short‐term changes in condition. We captured male and female goldfinches and measured bill color in the field and again under captive conditions several hours later. The following day, the captive birds were injected with either immunostimulatory lipopolysaccharide (LPS) or a control saline and changes in bill color were measured over a five day period. Yellow saturation of the bill decreased within 6.5 h between the field and captivity measures on the first day, presumably in response to capture stress. Over the longer experimental period, bill hue and luminance decreased significantly, whereas saturation significantly increased in both LPS and control groups. Bill hue and luminance decreased significantly more in birds treated with LPS than in control birds. Among LPS treated birds, individuals expressing high bill color at the beginning of the experiment lost more color than ‘low‐color’ birds, but still retained higher color at the end of the experiment, suggesting that birds that invest heavily in bill coloration are able to sustain high costs in the face of a challenge. Bill color change may have resulted from rapid reallocation of carotenoids from ornamentation to immune function. However, the complex shifts in bill color over time suggest that bill color may be influenced by multiple carotenoid compounds and/or changes in blood flow or chemistry in vessels just beneath the translucent keratinized outer layer of the bill. We conclude that bill color is a dynamic, condition‐dependent trait that strategically and reliably signals short‐term fluctuations in physiological condition.