Bill colour and immunocompetence in the European blackbird

The level of expression of secondary sexual characters has been suggested to signal male ability to resist parasitic infestations. To test this idea, several studies have examined the link between sexual signals and immunocompetence in birds. However, most of them have used only a single aspect of immune response to evaluate immunocompetence. We investigated the relation between bill colour and immunocompetence in captive male European blackbirds, Turdus merula, during the breeding season by assessing both cell-mediated and humoral components of the immune system. The blackbird is a sexually dimorphic species with bill colour varying from yellow to orange in males. Humoral immunity was assessed by measuring both primary and secondary responses to sheep red blood cell inoculation. Cell-mediated immunity was estimated with a delayed cutaneous hypersensitivity response to an injection of a mitogen (phytohaemagglutinin). No relation was found between male bill colour and the primary humoral response. However, males with orange bills showed a lower secondary humoral response but a higher cell-mediated immune response than males with yellow bills. Thus, the relation between immunocompetence and a secondary sexual trait may differ markedly depending on which component of the immune system is under consideration. We discuss our results in relation to mechanisms involved in sexual selection. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour      

Bill colour and correlates of male quality in blackbirds: an analysis using canonical ordination

Carotenoid-dependent plumage displays are widely assumed to be honest indicators of individual health or quality, which are used as cues during mate choice and/or agonistic signalling. Despite the fact that red, yellow and orange pigmentation of bills is common, and also variable between individuals, comparatively little is known about bill colouration as a condition-dependent trait. Furthermore, many studies of avian colouration are confounded by the lack of objective colour quantification and the use of overly simplistic univariate techniques for analysis of the relationship between the condition-dependent trait and individual quality variables. In this study, we correlated male blackbird bill colour (a likely carotenoid-dependent sexually selected trait) with body/condition variables that reflect male quality. We measured bill colour using photometric techniques, thus ensuring objectivity. The data were analysed using the multivariate statistical techniques of canonical ordination. Analyses based on reflectance spectra of male blackbird bill samples and colour components (i.e. hue, chroma and brightness) derived from the reflectance spectra were very similar. Analysing the entire reflectance spectra of blackbird bill samples with Redundancy Analysis (RDA) allowed examination of individual wavelengths and their specific associations with the body/condition variables. However, hue, chroma and brightness values also provided useful information to explain colour variation, and the two approaches may be complimentary. We did not find any significant associations between male blackbird bill colour and percent incidence of ectoparasites or cloaca size. However, both the colour component and full spectral analyses showed that culmen length explained a significant amount of variation in male blackbird bill colour. Culmen length was positively associated with greater reflectance from the bill samples at longer wavelengths and a higher hue value (i.e. more orange-pigmented bills). Larger males may have larger territories or be better at defending territories during male-male interactions, ensuring access to carotenoid food sources. Future studies should elucidate the relationship between bill colour and behavioural measures such as aggressiveness, territory size, song rate and nest attendance.     

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.     

The Role of Wing Pigmentation, UV and Fluorescence as Signals in a Neotropical Damselfly

Pigmentation patterns, ultraviolet reflection and fluorescent emission are often involved in mate recognition and mate quality functions in many animal taxa. We investigated the role of wing ultra-violet reflection, fluorescence emission, and pigmentation on age and sexual signals in the damselfly Mnesarete pudica. In this species, wings are sexually dimorphic in colour and exhibit age dependency: males and females show a smoky black colouration when young, turning red in mature males while it turns brown in females. First, we investigated wing UV patterns through reflectance and emission spectra. Second, behavioural experiments were undertaken to show male and female responses to manipulated wing pigmentation and experimentally reduced UV (UV-). Reflectance spectra of the wings of juvenile and mature males and females were used to show the differences between controls and individuals with manipulated colouration used in the behavioural experiment. UV-reduced, females with wings painted red, and control males and females were tethered and presented to conspecific males and females, and their behavioral responses were recorded. The male red wing pigmentation and females with red wings elicited an aggressive response in territorial males and a sexual response in females. Both males and females showed neutral responses towards individuals with reduced UV. Wing signals of juvenile individuals also provoked neutral responses. These results suggest that UV, together with pigmentation, plays a role during mate recognition in males and females. Other than butterflies and spiders, it seems that fluorescence signals and UV reflectance can also be part of communication in odonates.     

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.     

Sexual dichromatism in the yellow-breasted chat Icteria virens: spectrophotometric analysis and biochemical basis

Sexual dimorphism or dichromatism has long been considered the result of sexual selection. However, for many organisms the degree to which sexual dichromatism occurs has been determined within the confines of human perception. For birds, objective measures of plumage color have revealed previously unappreciated sexual dichromatism for several species. Here we present an unbiased assessment of plumage dichromatism in the yellow-breasted chat Icteria virens . Chats exhibit yellow to orange throat and breast plumage that to the unaided human observer differs only subtly in color. Spectrophotometric analyses revealed that chat throat and breast feathers exhibited reflective curves with two peaks, one in the ultraviolet and one in the yellow end of the spectrum. We found differences in both the shape and magnitude of reflectance curves between males and females. Moreover, for feathers collected from the lower edge and middle of the breast patch, male plumage reflected more light in the ultraviolet and yellow wavelengths compared to females, whereas male throat feathers appeared brighter than those of females only in the ultraviolet. Biochemical analyses indicated that the plumage pigmentation consisted solely of the carotenoid all- trans lutein and we found that males have higher concentrations of plumage carotenoids than females. Feathers that were naturally unpigmented reflected more UV light than yellow feathers, suggesting a potential role of feather microstructure in UV reflectance.     

Do Hosts Discriminate between Sexually Dichromatic Male and Female Brown‐headed Cowbirds?

Female brood parasites are recognized as threats to reproductive success by many host species. Male brood parasites may accompany females while they search for nests to parasitize and males depredate nests throughout the nesting cycle. Hence, selection may also favour recognition of males. We examined whether two common host species perceive male brown-headed cowbirds ( Molothrus ater ) as brood parasites, as nest predators, or neither. We quantified visits of male cowbirds to nests of yellow warblers ( Dendroica petechia ) and red-winged blackbirds ( Ageliaus phoeniceus ) to assess the frequency with which these host species interact with male cowbirds. Males were observed near nests during hosts' laying and incubating stages, although less frequently than female cowbirds. No visits by cowbirds occurred while parents cared for nestlings. We then presented models of male and female cowbirds plus a non-threatening control to yellow warblers and red-winged blackbirds during laying and nestling periods. If hosts perceive males and females similarly, they should respond more intensely to the cowbird models during the laying period, when nests are most likely to be parasitized. Both species responded similarly to male and female cowbird models during laying, which suggests that hosts view cowbirds of both sexes as threats. The responses of yellow warblers with nestlings to male cowbirds were strongly influenced by the order of model presentation. Warblers first presented with the male cowbird gave much reduced anti-parasite responses than those that first interacted with the female then the male cowbird. These results suggest that yellow warblers recognized male vs. female cowbirds, but that discrimination was not expressed during laying. By contrast, red-winged blackbirds did not discriminate between male and female cowbirds at either nesting stage.     

Discrimination among Males by Female Zebra Finches Based on Past as Well as Current Phenotype

There are many social and sexual contexts in which individuals should base their interactions with others on the recipient's history of behaviour or morphology over the time in which the two have associated. I tested the ability of female zebra finches to discriminate among males on the basis of their past as well as current phenotype by allowing them to view groups of four males for two hours a day, for four days, while the bill colour of individual males was manipulated with orange and red paint. Four bill colour treatments were assigned to the males in a stimulus set. One male was orange for all four days, one was red for all four days, a third male was orange for the first two days but red for the second two days, and the fourth male was red for the first two days and orange for the second two days. There was a slight, but non-significant, preference for orange bills when compared on a day to day basis regardless of bill colour history. However, when males were ranked on day 3 and day 4 according to the proportion of the assessment period they had been orange, there was a significant positive correlation between preference on those days and proportion of time orange. A more detailed examination shows that there was an effect of bill colour history independent of current bill colour but not vice-versa. These results suggest that discrimination is based not only upon instantaneous appearance, but average appearance over the assessment period.     

Wing pigmentation in territorial male damselflies, Calopteryx haemorrhoidalis: a possible relation to sexual selection

One striking characteristic in adult males of some odonate species is the presence of wing pigmentation. In Calopteryx species, males show a series of pre- and postcopulatory behavioural displays during which they face females while showing their pigmented wings. One hypothesis to explain the precopulatory flying displays and the associated wing pigmentation is that they may serve a sexual selection function. I investigated this in the territorial damselfly Calopteryx haemorrhoidalis. Males of this species defend aquatic substrates that females use for oviposition. Observational evidence indicated that males with a higher proportion of wing pigmentation were more likely to defend a territory, obtained more matings, had fewer gut parasites, survived in the study site and stayed in territories for longer. Experimental evidence suggested that the relationship mating success and wing pigmentation still held when controlling for the size of the substrate defended by territorial males. Similar to other studies in the Calopterygidae, these results suggest that wing pigmentation may be favoured by sexual selection. I discuss, however, whether an alternative function for male copulatory courtship displays and wing pigmentation, as sexual and/or species recognition, may also explain the evolution of these traits. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Evolution of sexual dimorphism in bill size and shape of hermit hummingbirds (Phaethornithinae): a role for ecological causation

Unambiguous examples of ecological causation of sexual dimorphism are rare, and the best evidence involves sexual differences in trophic morphology. We show that moderate female-biased sexual dimorphism in bill curvature is the ancestral condition in hermit hummingbirds (Phaethornithinae), and that it is greatly amplified in species such as Glaucis hirsutus and Phaethornis guy, where bills of females are 60 per cent more curved than bills of males. In contrast, bill curvature dimorphism is lost or reduced in a lineage of short-billed hermit species and in specialist Eutoxeres sicklebill hermits. In the hermits, males tend to be larger than females in the majority of species, although size dimorphism is typically small. Consistent with earlier studies of hummingbird feeding performance, both raw regressions of traits and phylogenetic independent contrasts supported the prediction that dimorphism in bill curvature of hermits is associated with longer bills. Some evidence indicates that differences between sexes of hermit hummingbirds are associated with differences in the use of food plants. We suggest that some hermit hummingbirds provide model organisms for studies of ecological causation of sexual dimorphism because their sexual dimorphism in bill curvature provides a diagnostic clue for the food plants that need to be monitored for studies of sexual differences in resource use.     

Female reproductive decisions and parasite burden in a calopterygid damselfly (Insecta: Odonata)

There is currently a gap in sexual selection theory about how much the environment drives female mating decisions. We present field data that suggest that female sexual behaviour in the damselfly Calopteryx haemorrhoidalis is influenced by parasite burden. Male wing pigmentation in Calopteryx is a sexually selected trait that signals a male's ability to cope with eugregarine parasites (an intestinal parasite that feeds on the adult's ingested food). Because adult C. haemorrhoidalis females also show wing pigmentation, we examined whether this trait is similarly influenced by parasite burden and whether it may signal the female's reproductive value. MaleC. haemorrhoidalis defend riverine substrates that females use for oviposition. After copulation and during oviposition, females are guarded by the copulating male against intruder males. Alternatively, females may avoid mating and ‘steal’ an oviposition site within a male's territory. In the present study, we found that the amount of female wing pigmentation was negatively correlated with the number of eugregarines present. Females with more parasites produced fewer eggs, survived fewer days, spent less time during courtship, ‘inspected’ fewer males before mating, had a lower mating success, were guarded for less time during oviposition and engaged in fewer ‘stealing’ events during oviposition. The reduced egg production and survival of heavily infected females may result from eugregarine depletion of the females' consumed food reserves. Thus, to offset reduced longevity, heavily infected females may accept a mating more rapidly and mate with fewer males. ‘Stealing’ behaviour may be related to the female's differential use of sperm from some males, particularly high-quality males. Interestingly, males that mated with low-pigmented females showed greater variance in wing pigmentation than did males that mated with high-pigmented females. Possibly, female wing pigmentation may signal a female's reproductive value, which provides females with longer mate-guarding episodes and reduced interference from intruder males. This study points out one possible constraint, intestine parasites, that females may face during mating decisions. Because females in bad condition mate with males in both good and bad condition, this constraint may be pervasive enough to weaken the intensity of selection for a male sexually selected trait, wing pigmentation, and help to maintain its variation in phenotypic expression. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Does beak coloration of male blackbirds play a role in intra and/or intersexual selection?

In many bird species, males may show brightly coloured traits and variance in male mating success may be explained by female preference and/or competition between males favouring the most coloured males. Male beak coloration has been suggested to play an important role in the pairing pattern of European blackbirds. Here, we investigate female preference and male-male interactions in relation to male beak coloration in this species. We used a field experiment to measure female and male responses toward stuffed decoys showing either of two beak coloration representing the extremes of the natural variation from yellow to orange. Decoys were situated on the centre of plots and behaviours of males and females approaching the decoy were recorded. The total number of males and females approaching the decoy did not differ between the two model treatments. Similarly, there was no difference in the mean time that males and females spent near the decoy. The number of males displaying aggressive postures did not differ between treatments and no female adopted a sexual posture. We discuss the relevance of our results and potential limitations associated with the experimental procedure.     

Female secondary sexual coloration and sex recognition in the keeled earless lizard, Holbrookia propinqua

Relationships between male social behaviour and female reproductive coloration in the iguanid lizard Holbrookia propinqua were examined by introduction of tethered non-resident lizards into the territories of adult males. Introduced lizards were plainly coloured females, females having bright reproductive coloration, adult males, plain females painted to resemble brightly coloured females, bright females painted to mimic plain females, and males painted with the bright yellow and orange secondary sexual coloration of females. Resident males courted all unpainted females, despite being aggressively rejected by the brightly coloured ones. They also courted all but two females in each of the painted groups. All unaltered non-resident males were challenged, attacked or subjected to other aggressive behaviour by residents, but all non-resident males painted to resemble bright females were courted. This differential treatment is highly significant. It clearly demonstrates that the bright female pigmentation functions in sex recognition, identifying a female to males. Other stimuli, especially pheromonal and behavioural cues, may contribute to sex recognition. Because sex recognition alone seems insufficient to account for the evolution of bright female coloration in H. propinqua and several other iguanids, several proposed additional functions are discussed.     

Trade-offs between immune investment and sexual signaling in male mallards

Allocation trade-offs between the immune system and sexual traits are central to current sexual selection hypotheses but remain contentious. Such trade-offs could be brought about by the dual action of testosterone that stimulates sexual signals but also suppresses immune functions and/or by competition for carotenoids that can be deposited in ornaments or used as antioxidants in support of immune functions. We investigated the trade-off between investment in immunity and maintenance of testosterone, carotenoids, and sexually selected, carotenoid-based bill color in male mallards. Following a nonpathogenic immune challenge, facultative immune investment resulted in a syndrome of changes in allocation. Plasma carotenoids disappeared from circulation proportional to antibody production. In addition, the reflectance spectrum of the bill was affected; greater antibody production was associated with an increase in relative UV reflectance. Although changes in bill reflectance and plasma carotenoids were related, the relationship appeared more complex than direct competition with immunity. Finally, maintenance of testosterone was affected by immune investment: testosterone levels declined substantially when males produced more antibodies. Because males with high testosterone are preferred by females, the decline in testosterone, in addition to carotenoid depletion and effects on bill reflectance, could constitute a significant cost of immune investment.     

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.     

Testosterone-Induced Expression of Male Colour Morphs in Females of the Polymorphic Tawny Dragon Lizard,Ctenophorus decresii

Many colour polymorphisms are present only in one sex, usually males, but proximate mechanisms controlling the expression of sex-limited colour polymorphisms have received little attention. Here, we test the hypothesis that artificial elevation of testosterone in females of the colour polymorphic tawny dragon lizard, Ctenophorus decresii, can induce them to express the same colour morphs, in similar frequencies, to those found in males. Male C. decresii, express four discrete throat colour morphs (orange, yellow, grey and an orange central patch surrounded by yellow). We used silastic implants to experimentally elevate testosterone levels in mature females to induce colour expression. Testosterone elevation resulted in a substantial increase in the proportion and intensity of orange but not yellow colouration, which was present in a subset of females prior to treatment. Consequently, females exhibited the same set of colour morphs as males, and we confirmed that these morphs are objectively classifiable, by using digital image analyses and spectral reflectance measurements, and occur in similar frequencies as in males. These results indicate that the influence of testosterone differs for different colours, suggesting that their expression may be governed by different proximate hormonal mechanisms. Thus, caution must be exercised when using artificial testosterone manipulation to induce female expression of sex-limited colour polymorphisms. Nevertheless, the ability to express sex-limited colours (in this case orange) to reveal the same, objectively classifiable morphs in similar frequencies to males suggests autosomal rather than sex-linked inheritance, and can facilitate further research on the genetic basis of colour polymorphism, including estimating heritability and selection on colour morphs from pedigree data.     

Ultraviolet reflectance by the cere of raptors

Ultraviolet (UV) signals have been shown to play key roles in social and sexual signalling in birds. Using a spectrophotometer, we analysed the colour of the cere (skin above the beak) of a diurnal raptor, the Montagu's harrier (Circus pygargus), and show that it reflects in the UV part of the spectrum. The cere is a well-known sexual signal in raptors, with carotenoid based pigmentation being indicative of quality. We thus hypothesized that UV reflectance also signals quality. Accordingly, we found that in our sample of wild males, the location of the UV peak was related to the orangeness of cere and correlated with male body mass and condition (mass corrected for size). Also, males with brighter UV were mated to females that laid earlier, as expected if UV reflectance relates to a male's quality and attractiveness. Future studies should investigate the relationships between UV reflectance and carotenoid pigmentation of cere, and test how UV reflectance influences mate choice.     

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.     

Experimental confirmation of the polygyny threshold model for red-winged blackbirds

The polygyny threshold model assumes that polygynous mating is costly to females and proposes that females pay the cost of polygyny only when compensated by obtaining a superior territory or male. We present, to the authors' knowledge, the first experimental field test to demonstrate that females trade mating status against territory quality as proposed by this hypothesis. Previous work has shown that female red-winged blackbirds (Agelaius phoeniceus) in Ontario prefer settling with unmated males and that this preference is adaptive because polygynous mating status lowers female reproductive success. Other evidence suggests that nesting over water increases the reproductive success of female red-winged blackbirds. Here we describe an experiment in which females were given choices between two adjacent territories, one owned by an unmated male without any over-water nesting sites and the other by an already-mated male with over-water sites. Females overwhelmingly preferred the already-mated males, demonstrating that superior territory quality can reverse preferences based on mating status and supporting the polygyny threshold model as the explanation for polygyny in this population.     

Carotenoid-based bill colour as an indicator of immunocompetence and sperm performance in male mallards

Female mate choice is often based on exaggerated sexual traits, signals of male qualities that females cannot assess directly. Two such key qualities are male immune and/or sexual competence, whereby honesty in signalling could be maintained by physiological trade-offs. Carotenoid-based ornaments likely constitute such honest signals, as there is direct competition for (limited) carotenoids between ornament deposition and anti-oxidant support of immune or sperm functioning. Using spectrometry, we assessed the potential signalling function of the yellow, carotenoid-based colour of the bill of male mallards, a target of female mate choice. Here we demonstrate that bill reflectance varied with plasma carotenoid level, indicating antioxidant reserves. Moreover, lower relative UV reflectance during autumn pairing predicted immune responsiveness and correlated positively with sperm velocity during breeding, a trait that affects fertility. Our data provide support for current theories that females could use carotenoid-based sexual signals to detect immune vigour and fertilizing ability of prospective mates.