Carotenoids bolster immunity during moult in a wild songbird with sexually selected plumage coloration

Carotenoid‐based colours in animals are valuable models for testing theories of sexual selection and life‐history trade‐offs because the pigments used in coloration are chemically tractable in the diet and in the body, where they serve multiple purposes (e.g. health enhancement, photoprotection). An important assumption underlying the hypothesized signalling value of carotenoid coloration is that there is a trade‐off in carotenoid pigment allocation, such that not all individuals can meet the physiological/morphological demands for carotenoids (i.e. carotenoids are limited) and that only those who have abundant supplies or fewer demands become the most colourful. Studies of carotenoid trade‐offs in colourful animals have been limited largely to domesticated species, which may have undergone artificial selection that changed the historical/natural immunomodulatory roles of carotenoids, to young animals lacking carotenoid‐based signals or to species displaying carotenoid‐based skin and bare parts. We studied the health benefits of carotenoids during moult in house finches (Carpodacus mexicanus), which display sexually selected, carotenoid‐based plumage coloration. We manipulated dietary carotenoid availability during both winter (nonmoult) and autumn (moult) in captive males and females and found that carotenoid‐supplemented birds mounted stronger immune responses (to phytohemagglutinin injection and to a bacterial inoculation in blood) than control birds only during moult. This study provides experimental, seasonal support for a fundamental tenet of Lozano's ‘carotenoid trade‐off’ hypothesis and adds to a growing list of animal species that benefit immunologically from ingesting higher dietary carotenoid levels. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 560–572.     

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.     

Testosterone treatment can increase circulating carotenoids but does not affect yellow carotenoid‐based plumage colour in blue tits Cyanistes caeruleus

A number of mechanisms are responsible for producing the variation in natural colours, and these need not act in isolation. A recent hypothesis states that carotenoid‐based coloration, in addition to carotenoid availability, is also enhanced by elevated levels of circulating testosterone (T). This has only been tested for carotenoid‐coloured bare parts in birds. We performed an experimental manipulation of T levels and examined the effects on the yellow carotenoid‐based breast plumage in captive yearling blue tits Cyanistes caeruleus, of which half received a diet supplemented with carotenoids. T treatment resulted in elevated plasma T compared to controls and carotenoid supplementation strongly increased plasma carotenoid levels. T treatment resulted in an additional increase in plasma carotenoid levels but only in the carotenoid‐supplemented males. Carotenoid supplementation resulted in more intense breast colour (carotenoid chroma), as expected. However, there was no effect of testosterone on plumage coloration at either dietary carotenoid level. Our results suggest that T can cause an increase in plasma carotenoid concentration, but that this does not necessarily lead to improved carotenoid‐based plumage coloration.     

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.     

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.     

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.     

Carotenoid‐based plumage coloration in golden‐crowned kinglets Regulus satrapa: pigment characterization and relationships with migratory timing and condition

Carotenoid‐based ornamental coloration has long been proposed to honestly signal quality due to its dependence on individual condition. Because migration can be one of the most stressful periods of an animal's annual cycle, developing colourful plumage may be particularly challenging for species in which migration and moult periods overlap or occur sequentially. The purpose of this study was to investigate pigmentary and condition‐dependent bases of carotenoid colour variation in a small migratory passerine, the golden‐crowned kinglet Regulus satrapa (Family Regulidae). We captured 186 male and female kinglets of various ages during fall migration in southwestern Ontario, Canada and recorded arrival date, body condition index, fat and pectoral muscle scores, wing mite infestation, and feather growth rate as measures of condition. We quantified crown coloration using reflectance spectrometry and analyzed feather carotenoids using high‐performance liquid chromatography. Yellow crown feathers of female kinglets contained only yellow hydroxycarotenoids, whereas orange feathers of males harboured a suite of eight carotenoid pigments. Males with longer wavelength orange crown hues deposited greater concentrations of ketocarotenoids, especially canthaxanthin. Female kinglets with longer wavelength crown hues and males with longer wavelength crown hues and more saturated crown coloration left for migration earlier in the year. Females with longer wavelength crown hues had fewer feather mites and tended to be in better condition. However, male kinglets with more saturated coloration possessed smaller pectoral muscles. This is the first study to identify plumage carotenoids in this North American bird family and to determine the pigmentary basis for both inter‐ and intrasexual colour variation. Our results provide further support for the condition‐dependence of carotenoid coloration and suggest that ornamental elaboration in both sexes may encode information about fall condition and migratory performance.     

Absorbance of retinal oil droplets of the budgerigar: sex,spatial and plumage morph-related variation

Intraspecific variation in photoreceptor physiology is known in several vertebrate taxa, but is currently unknown in birds, despite many avian traits varying intraspecifically, and avian visual ecology encompassing a wide range of environments and visual stimuli, which might influence spectral sensitivity. Avian retinal photoreceptors contain light absorbing carotenoid-rich oil droplets that affect vision. Carotenoids are also important plumage components. However, our understanding of the regulation of carotenoids in oil droplets remains rudimentary. Among birds, Melopsittacus undulatus has probably the best-studied colour vision, shows profound intraspecific variation in plumage colour, and increased plasma carotenoids during moult. We used microspectrophotometry to determine whether a relationship exists between oil droplet carotenoid concentration and plumage pigmentation, and tested for sex and spatial variation in droplet absorbance across the retina. Absorbance of one variety of P-type droplets was higher in males. No relationship was found between droplet absorbance and plumage colour. We found a spatial pattern of droplets absorbance across the retina that matched a pattern found in another parrot, and other avian species. Our work provides insights into the development and maintenance of retinal oil droplets and suggests a common mechanism and function for carotenoid deposition in the retina across bird species.     

Carotenoid‐based bill coloration functions as a social,not sexual,signal in songbirds (Aves: Passeriformes)

Many animals use coloration to communicate with other individuals. Although the signalling role of avian plumage colour is relatively well studied, there has been much less research on coloration in avian bare parts. However, bare parts could be highly informative signals as they can show rapid changes in coloration. We measured bill colour (a ubiquitous bare part) in over 1600 passerine species and tested whether interspecific variation in carotenoid‐based coloration is consistent with signalling to potential mates or signalling to potential rivals in a competitive context. Our results suggest that carotenoid bill coloration primarily evolved as a signal of dominance, as this type of coloration is more common in species that live in social groups in the nonbreeding season, and species that nest in colonies; two socio‐ecological conditions that promote frequent agonistic interactions with numerous and/or unfamiliar individuals. Additionally, our study suggests that carotenoid bill coloration is independent of the intensity of past sexual selection, as it is not related to either sexual dichromatism or sexual size dimorphism. These results pose a significant challenge to the conventional view that carotenoid‐based avian coloration has evolved as a developmentally costly, condition‐dependent sexual signal. We also suggest that bare part ornamentation may often signal different information than plumage ornaments.     

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.     

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.     

Season-, sex-, and age-specific accumulation of plasma carotenoid pigments in free-ranging white-winged crossbills Loxia leucoptera

Many birds acquire carotenoid pigments from foods and deposit these pigments into feathers and bare‐parts to become sexually attractive, but little work has been done on the interindividual and temporal variability in the types and amounts of carotenoids that free‐ranging individuals have available for use in coloration or other functions (e.g., in immunomodulation). To address this issue, we studied intra‐annual variation in plasma carotenoid profiles of juvenile and adult white‐winged crossbills Loxia leucoptera of both sexes. Adult male crossbills exhibit bright red carotenoid‐based plumage pigmentation, whereas females uniformly display drab yellow feather coloration and juvenile males only occasionally display some orange or pink color. Yellow xanthophylls (e.g., lutein, zeaxanthin) were predominant in plasma of birds from both sexes and age classes throughout the year. Plasma xanthophylls levels tended to be highest in the summer, when crossbills increase seed consumption for breeding as well as supplement their diet with insects. Blood accumulation of three other, less common plasma carotenoids‐β‐cryptoxanthin, rubixanthin, and gazaniaxanthin‐varied in a highly season‐, sex‐, and age‐dependent fashion. These carotenoids were virtually absent in juvenile or adult female plasma at all times of year and were only present in male plasma, at higher concentrations in adults than juveniles, during the period of feather growth (Sept.–Nov.). These pigments have been reported as valuable precursors of the metabolically derived red pigments (e.g., 3‐hydroxy‐echinenone, 4‐oxo‐rubixanthin, and 4‐oxo‐gazaniaxanthin, respectively) that appear in the plumage of male crossbills. These findings suggest that male crossbills either adopt a season‐specific foraging strategy to acquire foods rich in these pigments at the time they are needed to develop red coloration, or have a unique physiological ability to metabolically produce these pigments or absorb them from food during molt, in order to maximize color production.     

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.     

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.     

EVOLUTION OF CAROTENOID PIGMENTATION IN CACIQUES AND MEADOWLARKS (ICTERIDAE): REPEATED GAINS OF RED PLUMAGE COLORATION BY CAROTENOID C4‐OXYGENATION

Many animals use carotenoid pigments to produce yellow, orange, and red coloration. In birds, at least 10 carotenoid compounds have been documented in red feathers; most of these are produced through metabolic modification of dietary precursor compounds. However, it is poorly understood how lineages have evolved the biochemical mechanisms for producing red coloration. We used high‐performance liquid chromatography to identify the carotenoid compounds present in feathers from 15 species across two clades of blackbirds (the meadowlarks and allies, and the caciques and oropendolas; Icteridae), and mapped their presence or absence on a phylogeny. We found that the red plumage found in meadowlarks includes different carotenoid compounds than the red plumage found in caciques, indicating that these gains of red color are convergent. In contrast, we found that red coloration in two closely related lineages of caciques evolved twice by what appear to be similar biochemical mechanisms. The C4‐oxygenation of dietary carotenoids was responsible for each observed transition from yellow to red plumage coloration, and has been commonly reported by other researchers. This suggests that the C4‐oxygenation pathway may be a readily evolvable means to gain red coloration using carotenoids.     

Interspecific variation in the use of carotenoid-based coloration in birds: diet, life history and phylogeny

Birds show striking interspecific variation in their use of carotenoid-based coloration. Theory predicts that the use of carotenoids for coloration is closely associated with the availability of carotenoids in the diet but, although this prediction has been supported in single-species studies and those using small numbers of closely related species, there have been no broad-scale quantitative tests of the link between carotenoid coloration and diet. Here we test for such a link using modern comparative methods, a database on 140 families of birds and two alternative avian phylogenies. We show that carotenoid pigmentation is more common in the bare parts (legs, bill and skin) than in plumage, and that yellow coloration is more common than red. We also show that there is no simple, general association between the availability of carotenoids in the diet and the overall use of carotenoid-based coloration. However, when we look at plumage coloration separately from bare part coloration, we find there is a robust and significant association between diet and plumage coloration, but not between diet and bare part coloration. Similarly, when we look at yellow and red plumage colours separately, we find that the association between diet and coloration is typically stronger for red coloration than it is for yellow coloration. Finally, when we build multivariate models to explain variation in each type of carotenoid-based coloration we find that a variety of life history and ecological factors are associated with different aspects of coloration, with dietary carotenoids only being a consistent significant factor in the case of variation in plumage. All of these results remain qualitatively unchanged irrespective of the phylogeny used in the analyses, although in some cases the precise life history and ecological variables included in the multivariate models do vary. Taken together, these results indicate that the predicted link between carotenoid coloration and diet is idiosyncratic rather than general, being strongest with respect to plumage colours and weakest for bare part coloration. We therefore suggest that, although the carotenoid-based bird plumage may a good model for diet-mediated signalling, the use of carotenoids in bare part pigmentation may have a very different functional basis and may be more strongly influenced by genetic and physiological mechanisms, which currently remain relatively understudied.     

Carotenoid pigments in male American goldfinches: what is the optimal biochemical strategy for becoming colourful?

Studies of brilliant carotenoid‐based coloration in birds have traditionally centred on the role that these colours play in attracting mates. More recently, biologists have begun to take a biochemical approach to understanding the types of pigments found in feathers and how these relate to the expression of ornamental coloration. Nevertheless, surprisingly few studies have assessed the types and amounts of carotenoids present in the diet or blood of animals in relation to season, sex, condition or sexually attractive colour traits, particularly for wild birds. It is conceivable not only that the total concentration of pigments available is an important predictor of sexual attractiveness and mate quality, but also that specific pigments vary among individuals and play more important physiological and pigmenting roles than others. We investigated the carotenoid content of blood and feathers in wild‐caught, yellow‐pigmented male American goldfinches (Carduelis tristis) throughout the year to determine the optimal biochemical strategy for becoming colourful. We found that birds acquired two main yellow hydroxycarotenoids (lutein and zeaxanthin) from the diet during both moulting and non‐moulting periods. Blood concentrations of both pigments changed significantly over time, with moulting birds accumulating higher levels of both lutein and zeaxanthin, but proportionally more zeaxanthin, than non‐moulting birds. Moulting birds that acquired more lutein and more zeaxanthin in blood deposited a higher concentration of carotenoid pigments (canary xanthophylls A and B) into plumage and acquired more colourful feathers. In sum, these results indicate that (a) the types of dietary carotenoids available across seasons do not change in American goldfinches, (b) seasonal fluctuations in plasma‐carotenoid signatures may result from differences in dietary access or pigment processing, and (c) the best biochemical strategy for becoming a colourful, wild male goldfinch is to accumulate as many dietary/blood pigments as possible during moult. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 273–280.     

Plumage coloration can be perceived as a multiple condition‐dependent signal by Great Tits Parus major

The evolution of multiple signals can be explained because they enhance the perception of a general message by recipients. Plumage coloration frequently acts as a condition‐dependent signal, so that species displaying different colour patches have the potential to transmit information on condition through a multiple signalling system. The Great Tit Parus major exhibits plumage colour patches generated by the main types of colour production, some of which, particularly those based on melanins and carotenoids, are known to be related to body condition. However, the colour expressions of all of the different colour types have never been investigated simultaneously to determine whether they function as multiple signals of condition. In addition, visual perception models have never been applied to a multiple signalling system in a wild population of birds. Here I present information that links body condition with the colour expression of almost all of the different plumage patches of male Great Tits captured during the winter. Birds in better condition had greater reflectance values at short wavelengths in all plumage patches, and this was especially so in the white (i.e. structural) colour of the cheeks. Plumage colour characteristics were calculated by means of avian visual models, suggesting that Great Tits have the capacity to perceive information contained in the plumage coloration of conspecifics. These results show that short‐wavelength reflectance has great potential to transmit biologically significant information on the body condition of birds, even in achromatic plumage patches.     

Seasonal changes in carotenoid- and melanin-based plumage coloration in the Great Tit Parus major

Plumage coloration plays an important role as a signal of individual quality. We studied plumage coloration in two Great Tit Parus major populations to test the hypothesis that coloration changes during the year without moult. As expected, the coloration of 'yellow' and 'dark green' areas in the breast and back changed with time but not the colour characteristics of the 'black' crown. This suggests that colours based on carotenoids could be more exposed to abrasion or fading than melanin-derived colours, although other factors could affect the patterns of change found. Our results show that effects of colour alteration could be important to the understanding of the potential for reliable signalling of different colours. Seasonal changes in plumage colour should be considered in future analysis of intraspecific variation in coloration.     

Carotenoid‐based plumage colouration is predicted by age and parasites in the male European serin

A fundamental assumption of theories on the evolution of sexual signals is that they should be costly to produce in order to honestly signal the quality of the sender. The expression of carotenoid‐based plumage signals is considered to be condition‐dependent, due to the role of carotenoids functioning as pigments and as health modulators. We assessed carotenoid‐based plumage colouration in relation to male condition in a free living population of male European serins Serinus serinus during the breeding season. Male serins were trapped for morphometric and colouration measurements, during a four‐year field study, in order to evaluate the signalling value of colouration in relation to body condition and parasites level. We compared two different forms of colour quantification based on spectral data – the most commonly used tristimulus colour variables and physiological models of avian colour vision – and found that they were highly correlated for this species. We investigated the signalling value of male plumage colouration and it was found to be related to age and ectoparasite load. Plumage double cone and patch size were negatively related to parasites level, whereas SWS ratio was positively related to parasites and age. Colouration was also related with the time since moult. Our results indicate that the colour expression of serin's plumage is age dependent and is related, in complex ways, with the ability to cope with parasitic infection.