Dietary mineral content influences the expression of melanin-based ornamental coloration

Many animals develop bold patches of black or brown colorationthat are derived from melanin pigments and serve as sexual orsocial signals. At present, there is much debate among behavioralecologists over whether melanin-based color signals are costlyto produce. Studies that have manipulated crude aspects of nutrition(i.e., total food intake) or health have generally found melanin-basedplumage ornaments to be less responsive to such factors thanother types of extravagant color (e.g., carotenoid or structuralbased). However, a recently advanced hypothesis argues thatlimited minerals in the diet, such as calcium (Ca), zinc (Zn),and iron (Fe), may serve to increase melanin pigment productionand maintain signal honesty. Here, I experimentally tested whethervariation in the calcium content of the diet affects the colorand extent of melanin-based plumage in male zebra finches (Taeniopygiaguttata). Calcium supplementation increased the size, but notdarkness, of the black breast plumage patch in fledgling andadult males; however, sexually selected, carotenoid-based redbeak coloration was not affected by the diet manipulation. Theseresults are the first to support the idea that acquisition ofminerals from the diet is a unique, limiting factor for theexpression of ornamental melanin coloration in animals.     

Melanin-based plumage coloration in the house finch is unaffected by coccidial infection

For most species of birds, ornamental plumage coloration may result from two types of pigments: carotenoids and melanins. Despite the fact that melanin pigments can be synthesized by birds from basic, amino acid precursors, while carotenoids cannot be synthesized by birds and must be ingested, melanin-based plumage coloration and carotenoid-based plumage coloration have often been treated as a single trait in investigations of the function and evolution of plumage coloration. Expression of carotenoid-based coloration is known to be dependent on condition, while the effects of individual condition have not been well-tested for expression of melanin-based coloration. In this study, we experimentally tested the effect of coccidial infection of the intestinal tract of male house finches during moult on expression of melanin-based plumage coloration. Coccidial infection had a significant negative effect on carotenoid-based coloration, but it had no significant effect on melanin-based feather coloration. Unlike carotenoid-based coloration, melanin-based coloration may be cheap to produce, and honesty of melanin-based coloration my require social mediation.     

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.     

Corticosterone regulates multiple colour traits in Lacerta [Zootoca] vivipara males

Ornamental colours usually evolve as honest signals of quality, which is supported by the fact that they frequently depend on individual condition. It has generally been suggested that some, but not all types of ornamental colours are condition dependent, indicating that different evolutionary mechanisms underlie the evolution of multiple types of ornamental colours even when these are exhibited by the same species. Stress hormones, which negatively affect condition, have been shown to affect colour traits based on different pigments and structures, suggesting that they mediate condition dependence of multiple ornament types both among and within individuals. However, studies investigating effects of stress hormones on different ornament types within individuals are lacking, and thus, evidence for this hypothesis is scant. Here, we investigated whether corticosterone mediates condition dependence of multiple ornaments by manipulating corticosterone levels and body condition (via food availability) using a two‐factorial design and by assessing their effect on multiple colour traits in male common lizards. Corticosterone negatively affected ventral melanin‐ and carotenoid‐based coloration, whereas food availability did not affect coloration, despite its significant effect on body condition. The corticosterone effect on melanin‐ and carotenoid‐based coloration demonstrates the condition dependence of both ornaments. Moreover, corticosterone affected ventral coloration and had no effect on the nonsexually selected dorsal coloration, showing specific effects of corticosterone on ornamental ventral colours. This suggests that corticosterone simultaneously mediates condition dependence of multiple colour traits and that it therefore accounts for covariation among them, which may influence their evolution via correlational selection.     

Evolution of sexual dichromatism: contribution of carotenoid- versus melanin-based coloration

In birds, carotenoid-based plumage coloration is more dependent on physical condition and foraging abilities and less constrained developmentally than is melanin-based coloration. Thus, female mate choice for honest signals should result in more intense sexual selection on carotenoid- than on melanin-based plumage coloration. Using variation in sexual dimorphism as an indirect measure of the intensity of sexual selection, we tested the prediction mat variation in sexual dimorphism is driven more by change in carotenoid-based coloration between males and females dian by change in melanin-based coloration. Examination of historical changes in carotenoid- versus melanin-based pigmentation in 126 extant species of Cardueline finches supported this prediction. We found that carotenoid-derived coloration changed more frequendy among congeners dian melanin-based coloration. In both sexes, increase in carotenoid-based coloration score, but not in melanin-based coloration score, was strongly associated with increase in sexual dichromatism. In addition, sexual dimorphism in carotenoid-based coloration contributed more to overall dichromatism than dimorphism in melanin-based plumage. Our results supported die hypothesis that melanin-based and carotenoid-based coloration have fundamentally different signal content and suggest that combining melanin-based and carotenoid-based coloration in comparative analyses is not appropriate.     

Multivariate heredity of melanin-based coloration,body mass and immunity

The genetic covariation among different traits may cause the appearance of correlated response to selection on multivariate phenotypes. Genes responsible for the expression of melanin-based color traits are also involved in other important physiological functions such as immunity and metabolism by pleiotropy, suggesting the possibility of multivariate evolution. However, little is known about the relationship between melanin coloration and these functions at the additive genetic level in wild vertebrates. From a multivariate perspective, we simultaneously explored inheritance and selection of melanin coloration, body mass and phytohemagglutinin (PHA)-mediated immune response by using long-term data over an 18-year period collected in a wild population of the common kestrel Falco tinnunculus. Pedigree-based quantitative genetic analyses showed negative genetic covariance between melanin-based coloration and body mass in male adults and positive genetic covariance between body mass and PHA-mediated immune response in fledglings as predicted by pleiotropic effects of melanocortin receptor activity. Multiple selection analyses showed an increased fitness in male adults with intermediate phenotypic values for melanin color and body mass. In male fledglings, there was evidence for a disruptive selection on rump gray color, but a stabilizing selection on PHA-mediated immune response. Our results provide an insight into the evolution of multivariate traits genetically related with melanin-based coloration. The differences in multivariate inheritance and selection between male and female kestrels might have resulted in sexual dimorphism in size and color. When pleiotropic effects are present, coloration can evolve through a complex pathway involving correlated response to selection on multivariate traits.     

Sexual Dimorphism in Melanin Pigmentation,Feather Coloration and Its Heritability in the Barn Swallow (Hirundo rustica)

Melanin is the main pigment in animal coloration and considerable variation in the concentrations of the two melanin forms (pheo- and eumlanin) in pigmented tissues exists among populations and individuals. Melanin-based coloration is receiving increasing attention particularly in socio-sexual communication contexts because the melanocortin system has been hypothesized to provide a mechanistic basis for covariation between coloration and fitness traits. However, with few notable exceptions, little detailed information is available on inter-individual and inter-population variation in melanin pigmentation and on its environmental, genetic and ontogenetic components. Here, we investigate melanin-based coloration in an Italian population of a passerine bird, the barn swallow (Hirundo rustica rustica), its sex- and age-related variation, and heritability. The concentrations of eu- and pheomelanin in the throat (brown) and belly (white-to-brownish) feathers differed between sexes but not according to age. The relative concentration of either melanin (Pheo:Eu) differed between sexes in throat but not in belly feathers, and the concentrations in males compared to females were larger in belly than in throat feathers. There were weak correlations between the concentrations of melanins within as well as among plumage regions. Coloration of belly feathers was predicted by the concentration of both melanins whereas coloration of throat feathers was only predicted by pheomelanin in females. In addition, Pheo:Eu predicted coloration of throat feathers in females and that of belly feathers in males. Finally, we found high heritability of color of throat feathers. Melanization was found to differ from that recorded in Hirundo rustica rustica from Scotland or from H. r. erythrogaster from North America. Hence, present results show that pigmentation strategies vary in a complex manner according to sex and plumage region, and also among geographical populations, potentially reflecting adaptation to different natural and sexual selection regimes, and that some coloration components seem to be highly heritable.     

Melanin coloration in New World orioles I: carotenoid masking and pigment dichromatism in the orchard oriole complex

Carotenoids produce the brilliant red, orange, and yellow colors of many animals. However, melanin pigments can also confer some of these same hues. Because carotenoid and melanin colors are produced in different ways and may serve different signaling functions, either within or between species, it is important to establish whether one or both types of pigment are responsible for coloration. We have discovered what appears to be an evolutionary switch from carotenoid- to melanin-based color in two sexually dichromatic New World orioles. Using a combination of reflectance spectrometry and chromatographic analyses of plumage pigments, we found that the chestnut plumage of adult male orchard orioles Icterus spurius is produced predominantly by phaeomelanins. Orchard oriole feathers also contain carotenoids, which appear to be masked by the high concentration of phaeomelanins. In contrast, both carotenoids and phaeomelanins appear to contribute to color in adult male Fuertes's orioles I. fuertesi . Moreover, yellow yearling male and female plumage in both species is produced by carotenoids alone. The masking of carotenoids with phaeomelanins in orchard orioles is interesting in light of the signaling roles that carotenoids are thought to play. In addition, these plumage differences produce a unique case of age and sexual pigment dimorphism in orchard and Fuertes's orioles.     

Dietary carotenoids predict plumage coloration in wild house finches

Carotenoid pigments are a widespread source of ornamental coloration in vertebrates and expression of carotenoid-based colour displays has been shown to serve as an important criterion in female mate choice in birds and fishes. Unlike other integumentary pigments, carotenoids cannot be synthesized; they must be ingested. Carotenoid-based coloration is condition-dependent and has been shown to be affected by both parasites and nutritional condition. A controversial hypothesis is that the expression of carotenoid-based coloration in wild vertebrates is also affected by the amount and types of carotenoid pigments that are ingested. We tested this carotenoid-limitation hypothesis by sampling the gut contents of moulting house finches and comparing the concentration of carotenoid pigments in their gut contents with the colour of growing feathers. We found a positive association: males that ingested food with a higher concentration of carotenoid pigments grew brighter ornamental plumage. We also compared the concentration of carotenoids in the gut contents of males from two subspecies of house finches with small and large patches of carotenoid-based coloration. Consistent with the hypothesis that carotenoid access drives the evolution of carotenoid-based colour displays, males from the population with limited ornamentation had much lower concentrations of carotenoids in their gut contents than males from the population with extensive ornamentation. These observations support the idea that carotenoid intake plays a part in determining the plumage brightness of male house finches.     

Carotenoid-based ornamentation and status signaling in the house finch

The status signaling hypothesis (SSH) was devised primarilyto explain the adaptive significance of avian ornamental colorationduring the nonbreeding season. It proposes that conspicuousmale plumage serves as an honest signal of social status withina population of birds. However, to date this hypothesis hasbeen well tested and supported for only one type of plumage coloration, melanin-based coloration. Carotenoid-based pigmentationis known to positively reveal male health and condition duringmolt in a variety of species, but it is poorly understood whetherthis ornament type can also function as a status signal duringthe winter. We tested the SSH in male house finches (Carpodacusmexicanus) by manipulating the carotenoid-based plumage brightnessof first-year males and then pairing unfamiliar birds of differingcoloration in a series of dominance trials in captivity. Manipulated plumage color was unrelated to win/loss outcome in these trials.Similarly, the natural pigmentation of males was a poor predictorof winter dominance; as in other studies with this species,we found only a weak tendency for naturally drab males to dominatenaturally bright males. These results suggest that carotenoid-basedcoloration is not a reliable indicator of social status inmale house finches during the nonbreeding season. In fact, carotenoid-based coloration may function only in mate choice in this species,and it may be retained throughout the year either because timeconstraints preclude a second plumage molt or because it aidsin pair formation that begins in late winter.     

Energetic constraints on expression of carotenoid-based plumage coloration

Carotenoid pigments are used by many bird species as feather colorants, creating brilliant yellow, orange, and red plumage displays. Such carotenoid-based plumage coloration has been shown to function as an honest signal that is used in female mate choice. Despite recent interest in carotenoid-based ornamental traits, the basis for individual variation in expression of carotenoid-based plumage coloration remains incompletely understood. I tested the hypothesis that, independent of carotenoid access, food stress during molt would cause reduced expression of carotenoid pigmentation. I fed molting male House Finches Carpodacus mexicanus seed diets supplemented with either the red carotenoid pigment canthaxanthin or the yellow/orange carotenoid pigment β-cryptoxanthin (in the form of tangerine juice). Within each diet treatment, one group of males was given restricted food access and the other group was given unrestricted food access. Carotenoid supplements were placed in water so carotenoid access was controlled independent of food access. The results indicated a strong effect of both carotenoid access and food access on color display. Some males in the β-cryptoxanthin-supplemented group grew red plumage, suggesting that they can metabolically modify yellow pigments into red pigments, but no bird supplemented with β-cryptoxanthin grew plumage as red as birds supplemented with canthaxanthin. Males in the unrestricted food groups grew redder and more intensely pigmented plumage than males in the restricted food groups. These observations provide the best evidence to date of an energetic cost of carotenoid utilization in the generation of colorful plumage.     

Winter Plumage Coloration in Male American Goldfinches: Do Reduced Ornaments Serve Signaling Functions in the Non‐Breeding Season?

The signaling role of sexual ornaments that are displayed during the mating season is well known for many species, but dimorphisms that occur in the non‐breeding season have received much less attention, particularly when individuals only partially express their breeding condition during reproductively inactive periods. I assessed variation in the expression of colorful breeding and non‐breeding plumages in male American goldfinches (Carduelis tristis), a species in which males molt out of their colorful breeding ornaments in the fall but still display reduced carotenoid‐ and melanin‐based sexual dichromatism during the winter. I found that variability in the saturation of carotenoid‐based plumage pigmentation did not differ significantly between the breeding and non‐breeding seasons. Moreover, the area of melanin coloration in the cap was more variable in winter than when it is fully displayed during breeding. I also detected a significant positive correlation between the extent of melanin coloration during the winter and the saturation of non‐breeding carotenoid‐based plumage. Because of such variation in and correlated expression of these two color ornaments in winter, it is conceivable that male goldfinches display these hints of non‐breeding coloration for use as conspecific social or sexual signals. Natural selection pressures like predation and energetic demands are traditionally thought of as factors that restrict sex ornaments to breeding times alone, but this should not preclude animals from simply reducing their exaggerated features during winter and finding an expression optimum that balances signal costs and value. Such ‘remnant’ winter ornaments might be expected to evolve not only in animals that live in large non‐breeding groups (e.g. status‐signaling systems) but also in those where mates begin associating before breeding onset.     

Melanin coloration in New World orioles II: ancestral state reconstruction reveals lability in the use of carotenoids and phaeomelanins

Although many animals use carotenoids to produce bright yellow, orange, and red colors, an increasing number of studies have found that other pigments, such as melanins, may also be used to produce bright colors. Yet, almost nothing is known about the evolutionary history of this colorful melanin use. We used reflectance spectrometry to determine whether colors in New World orioles were predominantly due to carotenoids, colorful melanins, or a mixture of both. We then used ancestral state reconstruction to infer the directionality of any pigment changes and to test for phylogenetic signal. We found that three oriole taxa likely switched from carotenoid- to melanin-based colors. Several other oriole taxa apparently gained localized melanin coloration, or had coloration that seemed to be produced by a mixture of carotenoids and melanins. We also found little phylogenetic signal on the use of carotenoids or melanins to produce color. However, all pigment changes occurred within one of three major clades of the oriole genus, suggesting there may be signal at deeper phylogenetic levels. These repeated independent switches between carotenoid and melanin colors are surprising in light of the important signaling role that color pigments (especially carotenoids) are thought to play across a wide range of taxa.     

Carotenoid-Based Plumage Coloration Predicts Leukocyte Parameters during the Breeding Season in Northern Cardinals (Cardinalis cardinalis)

Dichromatism in songbirds is often associated with polygyny and dimorphic parental investment, and is thought to arise via sexual selection. Northern cardinals (Cardinalis cardinalis) are not only dichromatic but also monogamous and biparental, suggesting that plumage coloration in this species may serve different functions than in more typical dichromatic species. In order to explore the role of sexual selection in the evolution of plumage coloration in cardinals, we used reflectance spectrophotometry to investigate whether two carotenoid‐based ornaments, the male’s red breast and the female’s underwing coverts, contain information that potential mates or competitors could use to assess condition. We found that whereas coloration was not related to body condition (measured as the residual body mass from a regression of body mass on wing chord), more saturated carotenoid coloration was associated with higher heterophil to lymphocyte ratios in males, and with higher white blood cell counts in females. Thus, in both sexes, carotenoid coloration was positively linked to immune measures normally associated with higher levels of stress and infection. These results do not indicate that carotenoid‐based coloration functions as a signal of low levels of stress or disease in this species. We propose instead that because plumage coloration may be related to competitiveness, the more saturated individuals increase their risk of injury, stress, and infection by engaging in more competitive behavior or by secreting more testosterone, or both. Our finding that carotenoid pigmentation is positively associated in males with the size of the cloacal protuberance, an androgen‐sensitive sex character, supports this hypothesis.     

The role of pigment based plumage traits in resolving conflicts

The role of melanin ‘badges of status’, in male–male competition has been well‐studied, in contrast, carotenoid based plumage has largely been examined in the context of female mate choice. Recent work has shown that carotenoid signals can also function in male–male competition, although the functions of the two types of signals is currently unclear. Here, we examine the relationships between colouration, dominance and aggression in the crimson finch Neochmia phaeton, a species where males have both conspicuous red carotenoid plumage and a black melanin patch. We examined the importance of carotenoid and melanin based signals in three contexts: 1) among free‐living birds interacting at a feeding station: we found that neither colour signal influenced the outcome of interactions; 2) in staged dyadic contest in captivity: we found that coloration from carotenoid pigments was positively related to the probability of winning a contest, while the size of the melanin plumage patch was not related to winning; and 3) in staged dyadic contests where male plumage colour had been masked: we found that the number of interactions required to determine dominance increased. While the underlying natural plumage colour was still important in these contests, birds with more intense carotenoid colouration were now more likely to lose. These results confirm carotenoid‐based signalling in male–male contests. However this signal is used in conjunction with other factors such as self‐assessment and body condition. Contrary to traditional expectations, the black melanin patch was not found to be important in this context.     

The role of melanin- and carotenoid-based plumage coloration in nest defence in the Great Tit

Although plumage coloration is recognized to convey valuable information about the bearer's parental abilities, few studies have explored the relationship between coloration and nest defence. In this study in Great Tit Parus major, we analysed the relationship between nest defence and melanin‐ as well as carotenoid‐based plumage coloration, after controlling for ecological variables known to influence nest defence. A principal components analysis was applied to classify birds according to how vigorously they defended the nest, and the intensity of nest defence was tested against plumage coloration. Males with a large black tie defended their nests more vigorously, but no such effect was found for yellow breast coloration. This suggests that melanin‐based coloration in the Great Tit is associated with aggression, including both dominance‐aggression and nest defence, whereas carotenoid‐based coloration is not. The challenge in future studies will be to demonstrate whether females use this trait as an ornament to assess male quality and whether they trade off between the different ornaments a male may exhibit.     

Red-winged blackbirds Agelaius phoeniceus use carotenoid and melanin pigments to color their epaulets

Over the past three decades, the red‐winged blackbird Agelaius phoeniceus has served as a model species for studies of sexual selection and the evolution of ornamental traits. Particular attention has been paid to the role of the colorful red‐and‐yellow epaulets that are striking in males but reduced in females and juveniles. It has been assumed that carotenoid pigments bestow the brilliant red and yellow colors on epaulet feathers, but this has never been tested biochemically. Here, we use high‐performance liquid chromatography (HPLC) to describe the pigments present in these colorful feathers. Two red ketocarotenoids (astaxanthin and canthaxanthin) are responsible for the bright red hue of epaulets. Two yellow dietary precursors pigments (lutein and zeaxanthin) are also present in moderately high concentrations in red feathers. After extracting carotenoids, however, red feathers remained deep brown in color. HPLC tests show that melanin pigments (primarily eumelanin) are also found in the red‐pigmented barbules of epaulet feathers, at an approximately equal concentration to carotenoids. This appears to be an uncommon feature of carotenoid‐based ornamental plumage in birds, as was shown by comparable analyses of melanin in the yellow feathers of male American goldfinches Carduelis tristis and the red feathers of northern cardinals Cardinalis cardinalis, in which we detected virtually no melanins. Furthermore, the yellow bordering feathers of male epaulets are devoid of carotenoids (except when tinged with a carotenoid‐derived pink coloration on occasion) and instead are comprised of a high concentration of primarily phaeomelanin pigments. The dual pigment composition of red epaulet feathers and the melanin‐only basis for yellow coloration may have important implications for the honesty‐reinforcing mechanisms underlying ornamental epaulets in red‐winged blackbirds, and shed light on the difficulties researchers have had to date in characterizing the signaling function of this trait. As in several other birds, the melanic nature of feathers may explain why epaulets are used largely to settle aggressive contests rather than to attract mates.     

Melanin, carotenoid and structural plumage ornaments: information content and role in great tits Parus major

The importance of plumage colour as an indicator of individual quality and the basis of sexual selection has long been recognized. Of the three generally distinguished classes of plumage colours, melanin-based ornaments are traditionally considered to provide less reliable information than carotenoid-based traits. However, the role of structural ornaments in multiple signalling systems has rarely been examined, and no study has compared the information content and role of the three ornament types simultaneously. Here we investigated three plumage ornaments in great tits Parus major : the size of the melanin-based breast stripe, the carotenoid-based colour of the yellow breast and the structurally based reflectance properties of the black crown. We worked on both the mechanistic and the functional levels. First, we assessed the dependence of ornaments on body condition during moult using ptilochronology. Second, we estimated assortative mating for these traits, as a measure of mutual sexual selection. Only the spectral attributes of crown feathers correlated with body condition during moult. However, breast stripe size was related to age, while the brightness of the yellow breast indicated body size. Relative crown ultraviolet reflectance was much higher in males than in females. Assortative mating was strongest for crown ultraviolet reflectance, but composite measures suggest that a system of multiple sexually selected traits with different information content may work in this population. These data support the accumulating evidence that the condition-dependence of melanin and carotenoid coloration is not qualitatively different. They also suggest that more research should target the reflectance properties of dark plumage areas in general, and ultraviolet crown ornamentation in tits in particular.     

Effect of Dietary Carotenoid Supplementation on Food Intake and Immune Function in a Songbird with no Carotenoid Coloration

Studies of ornamental carotenoid coloration suggest that animals may have evolved specialized mechanisms for maximizing color expression and advertising their potential worth as a mate. For example, when given a choice of foods, many carotenoid‐pigmented fishes and birds select the more colorful, presumably carotenoid‐rich foods, and then accumulate these pigments at high levels in both the integument and systemically, in order to boost their immune system and hence directly advertise their health state with their colors. The majority of animals, however, do not exhibit sexually selected carotenoid coloration, which raises the question of whether they still optimize pigment intake and allocation in ways that boost endogenous accumulation and health. We tested the effect of carotenoid supplementation on food intake, carotenoid accumulation in blood, and both innate and adaptive immunity in male society finches (Lonchura domestica) – a non‐carotenoid‐colored estrildid finch relative of the zebra finch (Taeniopygia guttata; a species in which males do display sexually attractive and health‐revealing carotenoid color). Males fed a carotenoid‐rich diet for 2 wk did not consume more food than control males. Still, consumption of the carotenoid‐rich diet for 2 wk significantly elevated circulating levels of carotenoids in blood in male society finches, yielding the potential for immune enhancement. In fact, carotenoid‐enriched finches performed significantly better than control birds in our assay of constitutive innate immunity (bacterial‐killing activity of whole blood), although not in our test of inducible adaptive immunity (response to a mitogenic challenge with phytohemagglutinin). These results suggest that affinities for carotenoid‐rich foods may be particular to species with sexually selected carotenoid pigmentation, but that, as in humans and other mammals (e.g. mice, rats) without carotenoid color, the immune‐boosting action of carotenoids is conserved regardless of the strength of sexual selection on pigment use.     

Multiple behavioral contexts of a melanized tail display in a desert lizard

Conspicuous color displays in animals are often studied in the context of how they communicate an individual's reproductive value, ability to escape capture, or overall health. The study of sexual dichromatism often focuses on the evolution of male display traits balanced between detectability and mate acquisition, representing constraints stemming from both natural and sexual selection. Melanistic coloration and melanin color patterns play important roles in both intraspecific and interspecific signaling in animals; as melanin serves multiple purposes in organismal biology, the evolution of melanin-based visual displays might not be restricted to a single behavioral or ecological context. Here, we examine the behavioral contexts of the conspicuous melanin-based tail display in the greater earless lizard (Cophosaurus texanus), asking if this melanized tail pattern is associated with (a) female association preference of males varying in the concentration of melanin, (b) individual locomotor performance capacity, and (c) thermal ecology. We found the amount of tail melanin is linked with both locomotor performance and female association preference, with greater concentrations of melanin predicting the evolution of faster performance capacities and females preferring males with more discrete melanin bars. We found limited support that differences in melanin are linked with thermoregulation or thermal ecology. Overall, we find that multiple behavioral contexts shape the evolution of this melanized display trait. We conclude that behavioral constraints stemming from both natural and sexual selection can interact synergistically to favor the evolution and expression of a melanistic visual display, even in potentially costly ecological contexts.