Changes in carotenoid‐based plumage colour in relation to age in European Serins Serinus serinus

Yearling birds generally display duller colours than adults. This may be due to selection favouring birds with more intensely coloured plumage or to an increase in colour after the first complete moult. Most research to date on the topic has been carried out on species with structural plumage coloration or with carotenoid‐based coloration that is produced by the unmodified deposition of pigments. However, no study has been carried out on species whose carotenoids are metabolically modified before deposition. In this study, we assess age‐related changes in the carotenoid‐based coloration of European Serins, a species that metabolically processes carotenoids before they can be deposited into feathers. Birds were captured over consecutive years and we carried out both cross‐sectional and longitudinal analysis. Adults had significantly greater values of lightness and chroma than yearling birds. However, there were no changes in plumage colour when analysing the same individuals captured in subsequent seasons. Plumage lightness and chroma of adult males after moult were related to body mass, suggesting a role of body condition on plumage coloration. Our results suggest that changes in plumage coloration with age in European Serins are due to a selection process that favours more intensely coloured individuals.     

Nanostructure predicts intraspecific variation in ultraviolet-blue plumage colour

Evidence suggests that structural plumage colour can be an honest signal of individual quality, but the mechanisms responsible for the variation in expression of structural coloration within a species have not been identified. We used full-spectrum spectrometry and transmission electron microscopy to investigate the effect of variation in the nanostructure of the spongy layer on expression of structural ultraviolet (UV)-blue coloration in eastern bluebird (Sialia sialis) feathers. Fourier analysis revealed that feather nanostructure was highly organized but did not accurately predict variation in hue. Within the spongy layer of feather barbs, the number of circular keratin rods significantly predicted UV-violet chroma, whereas the standard error of the diameter of these rods significantly predicted spectral saturation. These observations show that the precision of nanostructural arrangement determines some colour variation in feathers.     

Plumage colour in nestling blue tits: sexual dichromatism,condition dependence and genetic effects

Sexual-selection theory assumes that there are costs associated with ornamental plumage coloration. While pigment-based ornaments have repeatedly been shown to be condition dependent, this has been more difficult to demonstrate for structural colours. We present evidence for condition dependence of both types of plumage colour in nestling blue tits (Parus caeruleus). Using reflectance spectrometry, we show that blue tit nestlings are sexually dichromatic, with males having more chromatic (more 'saturated') and ultraviolet (UV)-shifted tail coloration and more chromatic yellow breast coloration. The sexual dimorphism in nestling tail coloration is qualitatively similar to that of chick-feeding adults from the same population. By contrast, the breast plumage of adult birds is not sexually dichromatic in terms of chroma. In nestlings, the chroma of both tail and breast feathers is positively associated with condition (body mass on day 14). The UV/blue hue of the tail feathers is influenced by paternally inherited genes, as indicated by a maternal half-sibling comparison. We conclude that the expression of both carotenoid-based and structural coloration seems to be condition dependent in blue tit nestlings, and that there are additional genetic effects on the hue of the UV/blue tail feathers. The signalling or other functions of sexual dichromatism in nestlings remain obscure. Our study shows that nestling blue tits are suitable model organisms for the study of ontogenetic costs and heritability of both carotenoid-based and structural colour in birds.     

Carotenoid plumage hue and chroma signal different aspects of individual and habitat quality in tits

We hypothesized that Blue Tits Cyanistes caeruleus and Great Tits Parus major from low quality habitat (small woods) would have less yellow ventral plumage than those from high quality habitat (large woods) because they moult faster and/or their diet contains fewer carotenoids. They moult faster because they moult later in the season and are subject to more rapidly shortening daylengths. We tested this using a database of the plumage coloration (chroma, hue and lightness) of birds breeding in woods of different sizes, by manipulating the speed of moult in captive Blue Tits, and by counting the abundance and size of caterpillars (the major source of dietary carotenoids) in the diet of nestlings. In accordance with our hypothesis, juveniles of both species (which moult about three weeks later than adults) were about 8% less saturated in colour (lower chroma) than adults, but there was no significant difference in chroma between habitats. However, both species did differ significantly in hue between large and small woods. Blue Tits forced to moult faster in captivity, at a rate similar to that caused by a month's delay in the start of moult, had yellow flank feathers that were 32% less saturated in colour than those allowed to moult more slowly. Blue Tit nestlings in large woods consumed 47% more caterpillar flesh (per gram of faecal material voided) than those in small woods, and Great Tit pulli 81% more. When habitat effects were controlled for in ANOVAs, Blue Tits mated assortatively on the basis of flank hue and Great Tits on the basis of flank lightness. Flank colour therefore has the capacity to provide information about the potential quality of both habitats, and individual birds, to potential colonists and sexual partners.     

Sex and age differences in reflectance and biochemistry of carotenoid‐based colour variation in the great tit Parus major

The plumage coloration in great tits (Parus major) is the subject of much behavioural and ecophysiological research, yet there is a lack of analyses of the natural colour variation and its mechanisms. We used reflectance spectrometry and high‐performance liquid chromatography to explore individual, sexual and age‐related variation in carotenoid coloration and pigmentation, paramount to the often presumed, but rarely substantiated, costs and ‘honesty’ of carotenoid displays. In adults, we found that sex was the strongest predictor of ‘brightness’ (higher in males) and of ‘hue’ (longer wavelength in females). There was no sex difference in ‘carotenoid chroma’ or carotenoid content of feathers which also was unrelated to adult age (1 or 2+ years) and condition. Similar patterns were revealed for nestlings. Regarding the biochemical ‘signal content’, ‘carotenoid chroma’, but not ‘hue’, was significantly related to the carotenoid content (lutein and zeaxanthin) of feathers. These results refute the previously assumed exaggeration of carotenoid pigmentation in male great tits, and question the condition‐dependence of carotenoid coloration in this species. However, the sexual dimorphism in total reflectance or ‘brightness’, most likely due to melanins rather than carotenoids, may have implications for signalling or other adaptive explanations that need to be explored. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 758–765.     

Habitat structure and prey composition generate contrasting effects on carotenoid‐based coloration of great tit Parus major nestlings

Carotenoid‐based coloration of nestling plumage is generally considered a reliable signal of quality and has consistently been related to habitat structure. The main hypothesis proposed to explain this correlation is that high quality habitats contain high quality food, which in return affects the expression of carotenoid‐based plumage. It therefore assumes that, at the population level, the link between habitat structure and food composition is consistent and more relevant than inter‐individual differences in foraging ability or parental investment. In addition, it is assumed by default that food and habitat produce concordant effects on nestling coloration. In this work we evaluated habitat structure and prey composition in addition to several measures of parental investment. We investigated their relative effect on carotenoid‐based plumage coloration (lightness, chroma and hue) of great tit Parus major nestlings. We found a low correlation between carotenoid‐based coloration of nestlings and that of their parents. Nestling coloration, especially lightness and chroma, increased with the intake of more spiders. The time of breeding was positively correlated with lightness and chroma and negatively correlated with hue. Finally, the maturity of oak trees surrounding nest‐boxes correlated negatively with lightness, and the size of all tree species surrounding nest‐boxes correlated positively with hue of chick plumage. Our findings support the view that habitat structure and prey composition may produce divergent effects on feather pigmentation, and that prey proportions and variables related to parental investment should be assessed when considering carotenoid‐based coloration of chicks. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 547–555.     

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.     

The reliability of achromatic displays is island-dependent in nocturnal Storm Petrels

Nocturnal birds rely on achromatic visual signals to assess rivals and potential mates, but variation in the expression of these displays has been understudied. Here we use UV-visible reflectance spectrometry to study colour variation and the potential signalling function of the dark brown chest and white rump plumage – a colour pattern conspicuously exhibited during twilight courtship displays – in nocturnal Mediterranean Storm Petrels Hydrobates pelagicus melitensis from three islands with different exposures to predation and human disturbance. Human activity may increase perceived predation risk and thus affect activity budgets and the physiological stress response, with possible consequences on the relationship between individual body condition and plumage coloration. We found that chest and especially rump feathers reflected in the ultraviolet (UV) but there was no evidence of sexual dichromatism. However, we found support for a male-specific signalling role of plumage coloration. Indeed, chest yellow–red chroma and rump UV chroma were positively related to body condition in males, but only in the two islands representing the poorer environmental conditions. Therefore, environmental variability can potentially modify the reliability of achromatic displays, with possible consequences on sexual selection patterns in a nocturnal bird.     

The effects of diet quality and quantity on plumage colour and growth of great tit Parus major nestlings: a food manipulation experiment along a pollution gradient

The yellow carotenoid-based plumage coloration of great tit Parus major nestlings is found to be paler in polluted and urban environments. Because carotenoid pigmentation is often considered to be a condition dependent trait in birds we wanted to find out whether food-limitation and poor nestling condition could explain the pale plumage colour in a polluted area. P. major nestlings were supplemented with variable diets along a well known heavy metal pollution gradient around a copper smelter: two food treatments with carotenoids, one food treatment with little carotenoid and one unsupplemented control. Our field experiment showed that nestlings in the polluted area grew better with carotenoid rich diets, while such effect was not found in the unpolluted area. Nestlings showed higher plasma carotenoid (lutein) levels and higher plumage carotenoid chroma values in the unpolluted area than in the polluted area. However, plasma lutein levels or plumage colour were not associated with heavy metal levels in nestling faeces (a proxy for dietary exposure). Our results provide only weak evidence for carotenoid-based colouration to be condition-dependent in great tit nestlings as we found a positive relationship between body mass and carotenoid chroma in the non-supplemented control group only. The positive relationship between body mass and plumage colour intensity is more likely to be produced by the fact that good availability of caterpillars, an important food source for P. major, also means a good availability of carotenoids to nestlings. Our results suggest that main reason for pale nestling plumage in the polluted area is lower quality invertebrate food, and not nutrition-related oxidative stress.     

Cosmetic plumage coloration by iron oxides does not confer protection against feather wear

The active acquisition of plumage cosmetic coloration by iron oxides has been described in several bird species (notably the Bearded Vulture Gypaetus barbatus, but also in cranes, pelicans and other vulture species) but its function remains controversial. We tested the hypothesis that iron oxide acquired through mud bathing protects feathers from wear. We experimentally stained 13 breast feathers collected from two White Storks Ciconia ciconia with mud from a ferruginous pond regularly used by Bearded Vultures, while another 13 feathers (controls) were treated with water. All feathers were exposed to a controlled physical abrasion treatment. We found no differences in wear between control and stained feathers, which is counter to predictions of the wear protection hypothesis for the acquisition of this cosmetic plumage coloration.     

Preservation agents influence UV-coloration of plumage in museum bird skins

Plumage colour has always been a major criterion when describing and distinguishing bird taxa. Today, the use of reflection spectrophotometry is the most commonly used technique to study plumage coloration. A major advantage of this method is the opportunity of observing reflection beyond the human colour vision range—including the UV-waveband. Traditional taxonomic and phylogenetic research is often based on bird skins held in collections in natural history museums worldwide. Different agents for preservation have been used to prevent skins from being damaged by arthropod pests. Sometimes, parts of the plumage have been contaminated with stains from preservation agents. When dried, they are almost invisible to the human eye under normal sunlight conditions and cause no obvious change to feather coloration. However, some preservation agents contain fluorescent components which show up brightly when illuminated with UV-light. Furthermore, undetectable to the human eye, stains from these agents annihilate UV-reflection, preventing accurate data collection based on the UV-reflection of bird feathers. Measuring plumage parts which have been accidentally stained will lead to a relative underestimate of UV-reflection. In studying 20,000 samples, we found fluorescent stains in some 300 bird skins of varying ages (1913–2004) in museum collections throughout Europe and the USA. Different preservation agents have been evaluated for their fluorescence properties.     

Reduced sexual dichromatism,mutual ornamentation,and individual quality in the monogamous Zenaida dove Zenaida aurita

Although variation in plumage coloration is known to occur both between and within sexes, its study remains limited to a few bird families. The Zenaida dove Zenaida aurita is a socially monogamous tropical columbid bird species, characterized by an overall cinnamon‐brownish plumage and structural colorations on the head and neck. The species has been described as sexually dichromatic for plumage, although color differences between males and females are not obvious in the field. We investigated variation in the presumably melanin‐based color of the crown, mantle, breast, and belly, in the iridescent dark‐blue streaks on the head, and in the symmetric iridescent patches on the neck, over the whole spectrum visible to birds. Further, unlike most previous studies, we assessed covariation between plumage color and phenotypic traits in both males and females in relation to the putative signaling function of ornaments. Zenaida doves appeared to be slightly sexually dichromatic for the hue of pigment‐based colored areas, with males being on average more reddish than females. However, this difference was not discernible when considering the avian visual system. Conversely, although the reflectance spectra of iridescent plumage did not significantly differ between sexes in brightness, chroma or spectral position of the peaks, color discrimination analyses showed that individuals should be able to perceive between‐ or within‐sex differences in the color of the iridescent patch. In addition, several color parameters of brown and iridescent feathers were significantly related to territorial status, body condition, wing chord, and, albeit weakly, to individual multilocus heterozygosity. Overall, our results thus suggest that plumage color might be a reliable signal of quality in individuals of both sexes in this species. Further studies are needed to test the potential implication of plumage coloration in mate choice and mating patterns in the Zenaida dove.     

Carotenoid coloration in greenfinches is individually consistent irrespective of foraging ability

Carotenoid-based plumage coloration of birds has been hypothesized to honestly reflect individual quality, either because carotenoids are difficult to acquire via food or because of a trade-off in allocation of carotenoids between maintenance and signaling functions. We tested whether differential foraging ability is a necessary precondition for maintaining individual differences in carotenoid-based plumage coloration in male greenfinches (Carduelis chloris). Wild-caught birds were brought into captivity, where half of them were supplemented with carotenoids while the other half was maintained on a carotenoid-poor diet. Color of the yellow parts of tail feathers, grown under natural conditions, was compared with that of the replacement feathers, grown in captivity. Carotenoid supplementation increased feather chroma (saturation). Color of wild-grown feathers significantly correlated with the color of lab-grown feathers. This result demonstrates the existence of a significant component of variation in carotenoid coloration, which reflects physiological qualities or genetic differences among individuals independent of foraging ability. Among both experimental groups, plasma carotenoid concentration during feather growth strongly correlated with chroma of the feathers grown in captivity. This indicates that carotenoid-based plumage coloration can reveal circulating carotenoid levels over a very wide range of concentrations, suggesting the ample signaling potential of such a mechanism.     

Color expression in experimentally regrown feathers of an overwintering migratory bird: implications for signaling and seasonal interactions

Plumage coloration in birds plays a critical role in communication and can be under selection throughout the annual cycle as a sexual and social signal. However, for migratory birds, little is known about the acquisition and maintenance of colorful plumage during the nonbreeding period. Winter habitat could influence the quality of colorful plumage, ultimately carrying over to influence sexual selection and social interactions during the breeding period. In addition to the annual growth of colorful feathers, feather loss from agonistic interactions or predator avoidance could require birds to replace colorful feathers in winter or experience plumage degradation. We hypothesized that conditions on the wintering grounds of migratory birds influence the quality of colorful plumage. We predicted that the quality of American redstart (Setophaga ruticilla) tail feathers regrown after experimental removal in Jamaica, West Indies, would be positively associated with habitat quality, body condition, and testosterone. Both yearling (SY) and adult (ASY) males regrew feathers with lower red chroma, suggesting reduced carotenoid content. While we did not observe a change in hue in ASY males, SY males shifted from yellow to orange plumage resembling experimentally regrown ASY feathers. We did not observe any effects of habitat, testosterone, or mass change. Our results demonstrate that redstarts are limited in their ability to adequately replace colorful plumage, regardless of habitat, in winter. Thus, feather loss on the nonbreeding grounds can affect social signals, potentially negatively carrying over to the breeding period.     

Observations on skin colour changes in juvenile lumpsuckers

This study reports on an experimental investigation of colour change in young lumpsuckers Cyclopterus lumpus , employing Munsell colour charts to quantify skin hue, colour value and chroma. Juvenile lumpsuckers can live pelagically close to the sea surface, or can attach to floating seaweed and debris with their suckers. When swimming they adopt a cryptic default light green coloration, with the dorsum being darker than the belly (countershading). This default colour is also seen in complete darkness. When attached to objects they take a colour stimulus from below and to the sides and change colour. They can achieve good colour matches (noticeable in 3 min; complete within 15 min) with substrata that are green-yellow in colour, especially the brown alga Ascophyllum nodosum . Colour matching is largely dependent upon changes in colour value (i.e. lightness/darkness) produced by melanin dispersal or concentration in the melanophores. Thus, on artificial backgrounds outside the range of achievable hue matching, they still achieve good matchings of colour value. Slight hue changes were recorded (probably reflecting interaction of basic skin colour with melanin dispersal state), but chroma (brightness) variations were negligible. Lumpsucker colour depends upon light intensity, allowing sustained colour matching as weed illumination changes.     

Plumage yellowness predicts foraging ability in the blue tit Cyanistes caeruleus

Carotenoid‐based coloration in adult birds has been often regarded as an honest signal of individual quality. However, few studies have demonstrated a link between carotenoid display and the quantity or quality of resources provided to the offspring. The present study investigated the expression of a carotenoid‐based ornament, the breast plumage yellowness of the blue tit Cyanistes caeruleus, in relation to the level of parental provisioning effort and the amount of carotenoid‐rich prey provided to the young. The study was conducted in two forest types (evergreen and deciduous), which also allowed an exploration of the possible existence of habitat effects on the coloration of breeding birds. It was found that plumage colour intensity (carotenoid chroma) correlated positively with nestling provisioning rates of both males and females, supporting the good parent hypothesis. In addition, carotenoid chroma was positively related with the proportion of Lepidoptera larvae brought to the nest in both sexes. Female but not male coloration was positively linked to breeding success (proportion of fledged young). Nestling coloration did not correlate with that of their parents, nor the frequency with which they were fed. Hue and lightness of nestling's plumage correlated positively with body mass and tarsus length, respectively. The results obtained in the present study indicate that ventral plumage coloration in blue tits may advertise the ingested carotenoids (carotenoid foraging ability) and also their overall parental quality in terms of nestling provisioning rates. This suggests that plumage yellowness can be used as an indicator of foraging ability in this species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 418–429.     

Cryptic sexual dichromatism occurs across multiple types of plumage in the Green-backed Tit Parus monticolus

Several recent studies have found instances of cryptic sexual dichromatism within avian taxa. Although this dichromatism has been found in plumage produced through a variety of proximate mechanisms, little is known about how dichromatism varies across these types of plumage within a single species. We used a reflectance spectrometer to measure colour within the Green-backed Tit Parus monticolus , a species which displays multiple types of pigment and structural colours. We found significant differences in spectral measurements corresponding to hue, chroma, and brightness between male and female carotenoid, melanin, structural white, grey and structural blue plumage. The only plumage that did not appear to show sexual dichromatism was the olive plumage of the back. These findings suggest that the mechanism(s) producing cryptic dichromatism in the Green-backed Tit are non-specific and act across multiple types of plumage, rather than within a single type, such as carotenoid-based or structurally produced.     

Juvenile coloration of Florida Scrub-Jays ( Aphelocoma coerulescens ) is sexually dichromatic and correlated with condition

The Florida Scrub-Jay is a monogamous cooperative breeder in which both males and females display extensive structurally based blue plumage. Juveniles of this species exhibit blue tail and wing feathers that they begin growing as nestlings, and some of these feathers are retained throughout their first year. Although the birds appear to be sexually monochromatic, we assessed whether cryptic dichromatism exists in both the magnitude and pattern of coloration in tail feathers of juvenile Florida Scrub-Jays. We then determined whether variation in plumage coloration is associated with nutritional condition during molt. Tails of juvenile male Florida Scrub-Jays exhibit a greater proportion of UV reflectance than those of females. Mass at age 11 days and ptilochronology of the juvenile tail feathers were used as measures of individual nutritional condition during feather growth, and the latter was found to be positively associated with UV chroma. These data demonstrate that Florida Scrub-Jays are sexually dichromatic and suggest that variation in plumage color may be condition dependent, although we cannot rule out alternative explanations. Juvenile plumage coloration, therefore, has the potential to function as a signal of individual quality in both males and females.     

Preen waxes do not protect carotenoid plumage from bleaching by sunlight

The plumage coloration of wild birds often changes during the breeding season. One of the possible reasons for this is that sunlight, and particularly ultraviolet (UV) wavelengths, degrades the pigments responsible for plumage coloration. It has been suggested that birds may apply preen wax to feathers to protect feathers from bleaching. This hypothesis is tested by exposing carotenoid-based breast feathers of Great Tits to ambient light, light filtered to exclude UV and darkness. Preen waxes were experimentally removed from feather samples and the effect of light on coloration of treatment and control feathers compared. Ambient light had an effect on feather colour but preen wax did not. Feathers exposed to sun gradually became less saturated and hues shifted towards shorter wavelengths. This was not apparent in control feathers kept in darkness. Feathers exposed to full-spectra sunlight faded more than those that were kept in light with UV wavelengths removed. There was a decrease in brightness of feathers in both experimental and control groups, which was assumed to be an effect of dirt accumulation. This experiment confirmed earlier suspicions regarding the detrimental effects of UV irradiation on carotenoid-based coloration of avian feathers but failed to show any protective function of preen waxes. The possible consequences of these mechanisms of colour change for birds with regard to mating strategies are discussed.     

Effect of feather abrasion on structural coloration in male eastern bluebirds Sialia sialis

We used observations of male eastern bluebirds captured twice within a breeding season to test whether changes in structural coloration are related to feather abrasion. Between first and second broods, the UV chroma and brightness of feathers decreased, while hue shifted towards longer wavelengths. Observed changes were greatest for feathers on the head, least for feathers on the rump, and intermediate for feathers on the back. For head feathers, we found a significant correlation between reduction in barb length and UV chroma. Plumage coloration at first capture was correlated with change in UV chroma such that the most ornamented males tended to lose more coloration. Moreover, the magnitude of UV color change was positively related to the number of days between color measurements.To test whether these changes were caused by abrasive properties of the nesting sites, we randomly increased or decreased the abrasiveness of nesting‐box entrances by attaching sand paper or smooth plastic tape. The type of box entrance had no signicant effect on either coloration or barb length change. Our results suggest that feather abrasion is a factor in the seasonal color changes of bluebirds.