Sex-specific relationships between urbanization,parasitism,and plumage coloration in house finches

Historically,studies of condition-dependent signals in animals have been male-centric,but recent work suggests that female ornaments can also communicate indivi...     

Correlated changes in male plumage coloration and female mate choice in cardueline finches

The coevolution of extravagant male traits and female mate preferences is a central tenet of sexual selection theory. In lineages in which males have developed more elaborate sexual characters, females favour the most extreme expression of the trait. In some taxa, however, ornamental displays have evolved from more to less exaggerated states. Under these circumstances, it is unclear whether females show preferences for an ancestral male condition or for the current, less elaborate display. Here, we tested female mate preferences relative to male ornamental coloration in two species of cardueline finch (the American goldfinch, Carduelis tristis, and pine siskin, Carduelis pinus) that have evolved less elaborate carotenoid-based colour displays from more elaborately coloured ancestral states. We presented females of each species with a choice of males having either large patches of red colour (the elaborate, ancestral condition) or with species-typical patches of yellow colour (the less elaborate, derived state). Female goldfinches and siskins showed consistent preferences for the natural colour displays of males, and not for the more elaborate, ancestral colour pattern. Previous research on another cardueline finch taxon (a subspecies of the house finch, Carpodacus mexicanus griscomi), however, showed that females prefer more elaborate, ancestral coloration to the current form of reduced colour expression. The lack of congruence between male trait expression and female trait preference in the lineage with the most recently derived reduction in trait expression suggests that there may be evolutionary lags in the correspondence between male traits and female preferences. A shift in the expression of male coloration appears to be the first step towards the evolution of reduced colour displays in these finches.     

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.     

Geographic variation in the carotenoid plumage pigmentation of male house finches (Carpodacus mexicanus)

Geographic variation in both the colour and pattern of carotenoid plumage pigmentation displayed by males in two subspecies of house finches ( Carpodacus mexicanus frontalis and C. m. griscom ) was quantified. The extent of ventral carotenoid pigmentation (patch size) differed markedly between these two subspecies; frontalis males from the U.S. (New York, Michigan, California and Hawaii) displayed a medium patch extending from their throats to their lower bellies, while griscomi males sampled in Guerrero, Mexico displayed small patches restricted to their throats. Frontalis males sampled in Michigan and New York and griscomi males were relatively bright in colouration, while frontalis males sampled in Hawaii were relatively drab. Populations of frontalis in California showed substantial local variation in average male colouration: in two areas only 12 km apart males were as colourful and as drab as any population sampled. In aviary experiments in which they were fed either a plain seed diet or a diet supplemented with red carotenoid pigments during moult, males from all populations converged on a similar appearance, except that griscomi males attained a brighter plumage than frontalis males when their diet was supplemented with red pigments. Regardless of diet, the difference in patch size between frontalis and griscomi males persisted after moult in captivity. The author concludes that the difference in patch size between frontalis and griscomi males reflects genetic differences between these populations, but that the differences in the mean plumage colouration of males among populations reflect differences in the access that males have to carotenoid pigments during moult.     

Dietary carotenoid pigments and immune function in a songbird with extensive carotenoid-based plumage coloration

Carotenoid pigments can directly enhance the immune responsesof vertebrates, and they are used by many animals to createornamental color displays. It has been hypothesized that thesetwo functions of carotenoid pigments are linked: animals musttrade off use of carotenoid pigments for immune function versusornamental display. We tested two key predictions of this hypothesiswith captive American goldfinches, Carduelis tristis, a specieswith extensive carotenoid-based plumage coloration. First, wetested whether the immune systems of male goldfinches are carotenoidlimited during molt by supplying treatment groups with low,approximately normal, or high dietary access to lutein and zeaxanthin.Dietary treatment had a significant effect on plumage and billcolor but not on immunocompetence. We compared the cell-mediatedand humoral immune responses and the course of disease afterinfection for males in the different treatments. We observedno significant effect of the carotenoid content of diet on immuneresponse or disease resistance. Second, we tested whether therewas a positive relationship between immune function and expressionof ornamental coloration by comparing both the pre- and posttreatmentplumage coloration of males to their immune responses. We failedto find the predicted trade-off between ornament display andimmune function. These findings do not support the hypothesisthat songbirds with extensive carotenoid-based plumage displaystrade off the use of carotenoids for ornamentation versus immunefunction.     

An experimental test of the contributions and condition dependence of microstructure and carotenoids in yellow plumage coloration

A combination of structural and pigmentary components is responsible for many of the colour displays of animals. Despite the ubiquity of this type of coloration, neither the relative contribution of structures and pigments to variation in such colour displays nor the relative effects of extrinsic factors on the structural and pigment-based components of such colour has been determined. Understanding the sources of colour variation is important because structures and pigments may convey different information to conspecifics. In an experiment on captive American goldfinches Carduelis tristis, we manipulated two parameters, carotenoid availability and food availability, known to affect the expression of carotenoid pigments in a full-factorial design. Yellow feathers from these birds were then analysed in two ways. First, we used full-spectrum spectrometry and high-performance liquid chromatography to examine the extent to which variation in white structural colour and total carotenoid content was associated with variation in colour properties of feathers. The carotenoid content of yellow feathers predicted two colour parameters (principal component 1--representing high values of ultraviolet and yellow chroma and low values of violet-blue chroma-and hue). Two different colour parameters (violet-blue and yellow chroma) from white de-pigmented feathers, as well as carotenoid content, predicted reflectance measurements from yellow feathers. Second, we determined the relative effects of our experimental manipulations on white structural colour and yellow colour. Carotenoid availability directly affected yellow colour, while food availability affected it only in combination with carotenoid availability. None of our manipulations had significant effects on the expression of white structural colour. Our results suggest that the contribution of microstructures to variation in the expression of yellow coloration is less than the contribution of carotenoid content, and that carotenoid deposition is more dependent on extrinsic variability than is the production of white structural colour.     

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.     

Melanin-based plumage coloration and flight displays in plovers and allies

Plovers and their allies exhibit an impressive diversity of melanin-based plumage patterns ranging from non-melanized to completely melanized species. We use phylogenetic comparative methods to test whether melanization has evolved in relation to sexual selection for attracting mates, to selection for signalling territory defence, or to natural selection for camouflage. First, according to sexual-selection theory, melanized plumage has evolved to amplify the courtship displays of males. As predicted by this hypothesis, we found that males with aerial displays had more melanized plumage than males of ground-displaying species. In addition, sexual dimorphism in melanization was greater in species with display flights than in species with ground displays. Second, melanization may have evolved through social interactions to signal competitive ability in territory defence. We did not find evidence for this hypothesis, since breeding density was unrelated to the melanization of either sex. Finally, melanized plumage may camouflage the incubating parent. The latter hypothesis was not supported, since melanization was unrelated either to the darkness of nest substrate or the extent of vegetation cover. Taken together, our results are most consistent with the sexual-selection hypothesis, and suggest that melanized plumage has evolved to enhance the aerial displays of male plovers.     

Candidate genes for carotenoid coloration in vertebrates and their expression profiles in the carotenoid-containing plumage and bill of a wild bird

Carotenoid-based coloration has attracted much attention in evolutionary biology owing to its role in honest, condition-dependent signalling. Knowledge of the genetic pathways that regulate carotenoid coloration is crucial for an understanding of any trade-offs involved. We identified genes with potential roles in carotenoid coloration in vertebrates via (i) carotenoid uptake (SR-BI, CD36), (ii) binding and deposition (StAR1, MLN64, StAR4, StAR5, APOD, PLIN, GSTA2), and (iii) breakdown (BCO2, BCMO1). We examined the expression of these candidate loci in carotenoid-coloured tissues and several control tissues of the red-billed quelea (Quelea quelea), a species that exhibits a male breeding plumage colour polymorphism and sexually dimorphic variation in bill colour. All of the candidate genes except StAR1 were expressed in both the plumage and bill of queleas, indicating a potential role in carotenoid coloration in the quelea. However, no differences in the relative expression of any of the genes were found among the quelea carotenoid phenotypes, suggesting that other genes control the polymorphic and sexually dimorphic variation in carotenoid coloration observed in this species. Our identification of a number of potential carotenoid genes in different functional categories provides a critical starting point for future work on carotenoid colour regulation in vertebrate taxa.     

The significance of subadult plumage in Darwin's finches, Geospiza fortis

I studied delayed plumage maturation, a common phenomenon inpasserine birds, in a species of Darwin's finch, Geospiza fortis,on Isla Daphne Major, Galapagos, Ecuador. To address the questionof why males advertise their youth and immaturity with subadultplumage, I determined the fitness consequences of possessingdifferent plumage states at different ages. Rate of entry intoadult plumage is both variable and heritable. In two cohorts,individuals that took longer to enter full adult plumage surviveddry conditions significantly better than those that acquiredadult plumage quickly. Behavioral observations helped revealthe reasons for this higher survival. Advantages arose fromthe avoidance of conflict that occurred by signaling subordinatestatus with subadult plumage in both the nonbreeding and breedingseasons. Although blacker males had a significantly lower survival,those black males that did survive had a reproductive advantagein their first breeding season over males with less black intheir plumage from the same cohort and raised significantlymore offspring to fledging. Thus the study demonstrates selectionon the length of the subadult plumage stage. [Behav Ecol 1990;l:161–170]     

Carotenoid-based plumage coloration of male greenfinches reflects health and immunocompetence

Hypotheses of parasite-mediated sexual selection (PMSS) propose that elaborate male ornaments have evolved due to female preferences. Females would benefit from mating with more ornamental males if males' ornamentation signals their health status and ability to provide parasite resistance genes for the offspring. Carotenoid-based plumage coloration of birds has been hypothesised to honestly reflect an individual's health status due to trade-off in allocation of carotenoids between maintenance and signalling functions. The prediction of this hypothesis, namely that individuals with brighter plumage are able to mount stronger immune responses against novel antigens and reveal generally better health state, was tested in captive male greenfinches (Carduelis chloris). Greenfinches with brighter yellow breast feathers showed stronger humoral immune response against novel antigen (SRBC) while no relationship between plumage coloration and an estimate of cell-mediated immune responsiveness (PHA response) was detected. Elaborately ornamental individuals had better general health state as indicated by the negative correlations between plumage brightness and heterophil haemoconcentration. Consistent with the concept of PMSS, these results suggest that carotenoid-based plumage coloration in greenfinches honestly signals immunocompetence and health status.     

Geographic variation in the plumage coloration of willow flycatchers Empidonax traillii

The ability to identify distinct taxonomic groups of birds (species, subspecies, geographic races) can advance ecological research efforts by determining connectivity between the non‐breeding and breeding grounds for migrant species, identifying the origin of migrants, and helping to refine boundaries between subspecies or geographic races. Multiple methods are available to identify taxonomic groups (e.g., morphology, genetics), and one that has played an important role for avian taxonomists over the years is plumage coloration. With the advent of electronic devices that can quickly and accurately quantify plumage coloration, the potential of using coloration as an identifier for distinct taxonomic groups, even when differences are subtle, becomes possible. In this study, we evaluated the degree to which plumage coloration differs among the four subspecies of the willow flycatcher Empidonax traillii, evaluated sources of variation, and considered the utility of plumage coloration to assign subspecies membership for individuals of unknown origin. We used a colorimeter to measure plumage coloration of 374 adult willow flycatchers from 29 locations across their breeding range in 2004 and 2005. We found strong statistical differences among the mean plumage coloration values of the four subspecies; however, while individuals tended to group around their respective subspecies’ mean color value, the dispersion of individuals around such means overlapped. Mean color values for each breeding site of the three western subspecies clustered together, but the eastern subspecies’ color values were dispersed among the other subspecies, rather than distinctly clustered. Additionally, sites along boundaries showed evidence of intergradation and intermediate coloration patterns. We evaluated the predictive power of colorimeter measurements on flycatchers by constructing a canonical discriminant model to predict subspecies origin of migrants passing through the southwestern U.S. Considering only western subspecies, we found that individuals can be assigned with reasonable certainty. Applying the model to migrants sampled along the Colorado River in Mexico and the U.S. suggests different migration patterns for the three western subspecies. We believe that the use of plumage coloration, as measured by electronic devices, can provide a powerful tool to look at ecological questions in a wide range of avian species.     

The adaptive function of melanin-based plumage coloration to trace metals

Trace metals produced by anthropogenic activities are of major importance in urban areas and might constitute a new evolutionary force selecting for the ability to cope with their deleterious effects. Interestingly, melanin pigments are known to bind metal ions, thereby potentially sequestering them in inert body parts such as coat and feathers, and facilitating body detoxification. Thus, a more melanic plumage or coat coloration could bring a selective advantage for animals living in polluted areas. We tested this hypothesis by investigating the link between melanin-based coloration and zinc and lead concentrations in feathers of urban feral pigeons, both at capture time and after one year of captivity in standardized conditions. Results show that differently coloured pigeons had similar metal concentrations at capture time. Metal concentrations strongly decreased after one year in standardized conditions, and more melanic pigeons had higher concentrations of zinc (but not lead) in their feathers. This suggests that more melanic pigeons have a higher ability to store some metals in their feathers compared with their paler counterparts, which could explain their higher success in urbanized areas. Overall, this work suggests that trace metal pollution may exert new selective forces favouring more melanic phenotypes in polluted environments.     

Delayed maturation of plumage coloration and plumage spottedness in the Barn Owl (Tyto alba)

Summary The Barn Owl (Tyto alba) varies in plumage from dark reddish-brown to white, and from heavily marked with black spots to immaculate. Males are commonly lighter coloured and less spotted than females. I assessed whether male and female Barn Owls delay the full expression of plumage coloration and spottedness to the second year of life. In Switzerland, I quantified the two traits of birds captured at the nestling stage, first, second and third year of life. Males and females became lighter coloured only from the first to the second year. Males became less spotted only from the first to the second year, and females less spotted from the nestling stage to the first year but more spotted from the first to the second year. Females were also similarly spotted at the second and third year of age. By cutting off small pieces of feathers of females I could recognize which feathers had later been renewed. After a complete moult old females did not change in plumage characteristics.

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Verzögerter Wechsel vom Juvenil- zum Adultgefieder bei der Schleiereule(Tyto alba)

Zusammenfassung Das Gefieder der Schleiereule (Tyto alba) variiert von rostbraun bis weiss und von dicht gefleckt bis fleckenlos. Die Männchen sind eher hell und weniger gefleckt als die Weibchen. Es wurde untersucht, ob männliche und weibliche Schleiereulen die volle Entfaltung ihrer Gefiederfarbe und -struktur auf das zweite Lebensjahr verlegen. In der Schweiz habe ich beide Gefiederpolymorphismen bei Jungvögeln, ein-, zwei- und dreijährigen Vögeln quantifiziert. Bei Männchen und Weibchen wurde das Auslichten der Gefiederfarbe nur zwischen dem 1. und 2. Lebensjahr beobachtet. Männchen wurden weniger dicht gefleckt nur zwischen dem 1. und 2. Lebensjahr, während Weibchen zwischen dem Nestlingstadium und dem 1. Lebensjahr weniger gefleckt wurden und zwischen dem 1. und 2. Lebensjahr wieder mehr Flecken hatten. Weibchen wurden also gleich gefleckt im 2. und 3. Lebensjahr. Um die erneuerten Federn zu erkennen, wurden bei Weibchen kleine Federstücke herausgeschnitten. Nach einer vollständigen Erneuerung der Brustfedern hatte sich die Farbe und die Anzahl Punkte bei alten Weibchen nicht geändert.

     

Condition-dependent variation in the blue-ultraviolet coloration of a structurally based plumage ornament

After years of investigation into the function of sexually dimorphic ornamental traits, researchers are beginning to understand how bright plumage colour in birds acts as an intraspecific signal. This work has focused primarily on pigment-based ornaments because they are highly variable in patch size, hue and brightness for some species. In contrast, structurally based ornaments have been little studied, in part because they do not appear to be as variable as pigment-based ornaments. We investigated a structurally based plumage ornament in a wild population of blue grosbeaks (Guiraca caerulea), a sexually dimorphic passerine. We report plumage variation that extends into the ultraviolet region of the spectrum. The pattern of covariation between four out of five elements of plumage variation suggests that structurally based ornamentation is pushed towards extreme expression of the trait as predicted by the sexual selection theory. The ''bluest'' birds have the highest percentage of blue feathers on the body. These ornamental feathers reflect light maximally at the shortest wavelengths (ultraviolet), with the greatest intensity and the greatest contrast. Age may have some effect on expression of blueness. In addition, plumage variables are correlated with growth bars in tail feathers (a record of nutritional condition during moult in a non-ornamental trait). This suggests that the ornament is partially condition dependent. Thus, blue plumage in male grosbeaks may serve as an honest indicator of age and quality.     

The evolution of carotenoid coloration in estrildid finches: a biochemical analysis

The estrildid finches (Aves: Passeriformes: Estrildidae) of Africa, Asia, and Australia have been the focus of several recent tests of sexual selection theory. Many estrildids display bright red, orange, or yellow colors in the beak or plumage, which typically are generated by the presence of carotenoid pigments. In this study, we used high-performance liquid chromatography to investigate the carotenoid content of feathers and other colorful tissues in seven species of estrildids. Star finches (Neochmia ruficauda) and diamond firetails (Stagonopleura guttata) circulated two main dietary carotenoids (lutein and zeaxanthin) through the blood and liver and used both to acquire a yellow plumage color. However, five other estrildids (common waxbill, Estrilda astrild; black-rumped waxbill, Estrilda troglodytes; zebra waxbill, Amandava subflava; red avadavat, Amandava amandava; and zebra finch, Taeniopygia guttata) circulated these same dietary carotenoids along with two metabolites (dehydrolutein and anhydrolutein) through the blood and/or liver and used all four as yellow plumage colorants. We subsequently tracked the distribution of these pigments using a published phylogeny of estrildid finches to determine the evolutionary pattern of carotenoid metabolism in these birds. We found that finches from the most ancient tribe of estrildids (Estrildini) possessed the ability to metabolize dietary carotenoids. Although carotenoids from the most ancestral extant estrildid species have yet to be analyzed, we hypothesize (based on their relationships with other songbirds known to have such metabolic capabilities) that these finches inherited from their ancestors the capability to metabolize carotenoids. Interestingly, later in estrildid evolution, certain taxa lost the ability to metabolize dietary carotenoids (e.g., in the Poephilini), suggesting that the occurrence of carotenoid metabolism can be labile and is likely shaped by the relative costs and benefits of color signaling across different species.     

The effect of capture-and-handling stress on carotenoid-based beak coloration in zebra finches

Stress can have widespread effects on animal behaviors and phenotypes, including sexually selected traits. Ornamental colors have long been studied as honest signals of condition, but few studies have been conducted on how the physiological stress response (i.e., corticosterone (CORT) elevation) impacts color expression. We used a traditional capture-and-restraint technique to examine the effect of repeated handling stress on carotenoid-dependent beak coloration in male and female zebra finches (Taeniopygia guttata). Birds subjected to daily, 10-min handling treatments, which elevated circulating CORT levels, for a four-week period displayed deeper orange/red beak coloration than did control animals. Stressed males lost body mass during the experiment and marginally decreased in circulating carotenoid concentrations. Hence, handling stress may have reduced food intake or induced mobilization of body stores (i.e., fat) of carotenoids. In contrast to males, stressed females maintained orange beak color, while control females faded in color. This study highlights sex- and pigment-specific mechanisms by which stress may temporarily enhance the expression of sexual traits, but at the expense of other key fitness traits (e.g., body mass maintenance, reproduction).