Carotenoid-based Plumage Coloration Research Articles

Diet and between-tissue isotope comparisons reveal different foraging strategies for age and sex of a Saffron Finch (Sicalis flaveola Linnaeus, 1766) population.

Measuring stable isotopes in different tissues offers the opportunity to provide insight into the foraging ecology of a species. This study aimed to assess how diet varies between yellow females, yellow males, and dull individuals of a Saffron Finch (Sicalis flaveola) population. We measured δ13C and δ15N in blood over a year, and in different feathers, to estimate seasonal consistency of resource use for each category. We conducted this study in a private farm in the Central Brazilian savannas. We sampled 195 individuals in seven field samplings between January 2017 and March 2018. The mean blood δ13C values were similar among yellow females, yellow males and dull individuals, indicating that this population of Saffron Finch predominantly accesses similar resources throughout the year, with a predominant C4 signal. Although Saffron Finch is considered a granivorous species, the mean δ15N values found indicate that both adults and juveniles also incorporate in their tissues some invertebrate. The slight isotope-tissue difference between feathers and blood is similar to the reported in previous studies and may reflect tissue-to-tissue discrimination. The isotopic space of yellow males was greater than that of yellow females and dull individuals, indicating greater dietary diversity due to greater inter-individual variation in diet. In Saffron Finch, which delays plumage maturation, competition-driven partitioning of food resources seems essential in driving carotenoid-based plumage coloration between age classes and sexes.

Multiple ornaments: trade-offs and redundancy in signalling functions in male and female varied tits (Sittiparus varius)

Abstract Signals conveyed by ornaments can be indicators in the process of sexual selection. Specifically, signals may be superimposed on each other to represent an individual’s quality, convey different information, or offer redundant information. As much of the research in this area has focused on relationships between diverse signals in males, those of females have largely been overlooked. From 2022 to 2023, we tested whether three traits, namely acoustic (song display) and visual ornaments (patch size and carotenoid-based plumage coloration), and morphology (wing and tail length), were interrelated in males and whether visual ornaments and morphology were interrelated in females in a wild population of varied tits (Sittiparus varius). We found that males with brighter and more reddish plumage sang at lower maximum and minimum frequencies, with narrower bandwidths and lower frequency peaks. Similarly, a negative correlation between tail length and carotenoid-based plumage coloration was observed in both sexes. Furthermore, a positive relationship was found between signals within traits. These results highlight the trade-offs in signals between different traits, as well as the consistency within the same traits.

Phenotypic associations of common waxbill gut and feather microbiome diversity in a shared environment

Abstract Microbial diversity may affect host phenotypes, including morphology and behaviour, but it remains unclear to which extent those could be mediated by host differences in diet or habitat use. We used a population of wild common waxbills living in a large open-air mesocosm to control for such differences and test whether phenotypic variation is associated with cloaca bacteria diversity. We also tested correlations between feather-associated bacteria diversity and plumage colour ornamentation. Albeit weakly correlated, individuals sharing more similar cloaca microbiotas were more similar in their carotenoid-based plumage and bill red colour, suggesting that the gut microbiota can influence ornamentation even when individuals are in the same habitat and have access to the same food resources, perhaps mediating individual differences in feeding or digestion. There were no associations between feather bacteria and plumage colour, nor between cloaca bacteria and body size or several behavioural phenotypes. These results further highlight that investigating associations between microbiota and behaviour in nature should account for variation in extrinsic factors.

Chromatic and achromatic differences of melanin- and carotenoid-based plumage coloration in five minivet species (Pericrocotus spp.) under conspecific and predator visual systems

Adaptive mate choice has been accepted as the leading theory to explain the colorful plumage of birds. This theory hypothesizes that conspicuous colors act as signals to advertise the qualities of the owners. However, a dilemma arises in that conspicuous colors may not only attract mates, but also alert predators. The “private channels of communication” hypothesis proposes that some intraspecific signals may not be visible to heterospecific animals because of different visual systems. To better understand the evolution of plumage colors and sexual selection in birds, here we studied the chromatic difference and achromatic differences of melanin- and carotenoid-based plumage coloration in five minivet species (Pericrocotus spp.) under conspecific and predator visual systems. We found that either the chromatic or achromatic difference among male or female minivets’ plumage was consistently higher under conspecific vision than under predator vision for all five studied species of minivets. This result indicated that individual differences in plumage colors of minivets were visible to the conspecific receivers and hidden from potential predators as a result of evolution under predation risk and conspecific communication. However, males were under a higher risk of predation because they were more conspicuous than females to the vision of a nocturnal predator.

No evidence that endohelminth parasites cause selection against hybrid orioles across the Baltimore–Bullock’s Oriole hybrid zone

Abstract The Baltimore–Bullock’s Oriole hybrid zone is one of the best-studied avian hybrid zones in North America, yet our understanding of the causes of selection against hybrids remains poor. We examine if endohelminth parasites may cause selection against hybrid orioles but found no evidence for this hypothesis. Of the 139 male orioles we examined, 43 individuals contained endohelminth parasites from at least 1 of these groups: Cestoda, Acanthocephala, or Nematoda. Across the hybrid zone, Baltimore Orioles (Icterus galbula) and Bullock’s Orioles (I. bullockii) differed in their parasite communities, such that Baltimore Orioles frequently contained both Acanthocephala and Cestoda parasites whereas Bullock’s Orioles primarily contained Cestoda parasites. Despite these differences in parasite communities between parental species, the frequency of hybrid orioles with parasites was similar to parentals, suggesting that hybrids were as susceptible to endohelminth parasites as parentals. Using a subset of 99 adult male orioles, we explored how parasites may be associated with the expression of orange carotenoid-based plumage in hybrids and parentals. Associations between carotenoid-based plumage color and parasites were most strongly expressed in Bullock’s Orioles, but patterns were subtle and counterintuitive because individuals with parasites often had more enhanced color measures compared to individuals without parasites. Taken together, these data suggest that endohelminth parasites impose little fitness costs to male orioles on the breeding grounds and likely do not cause selection against hybrids.

Bigger or long-winged male common crossbills exhibit redder carotenoid-based plumage coloration.

Carotenoid-based ornaments are often considered reliable (honest) individual condition signals because their expression implies physiological costs unaffordable for low-quality animals (handicap signals). Recently, it has been suggested that efficient cell respiration is mandatory for producing red ketocarotenoids from dietary yellow carotenoids. This implies that red colorations should be entirely unfalsifiable and independent of expression costs (index signals). In a precedent study, male common crossbills, Loxia curvirostra, showing a red plumage reported higher apparent survival than those showing yellowish-orange colors. The plumage redness in this species is due to ketocarotenoid accumulation in feathers. Here, we correlated the male plumage redness (a 4-level visual score: yellow, patchy, orange, and red) and the body morphology in more than 1,000 adult crossbills captured in 3 Iberian localities to infer the mechanisms responsible for color evolution. A principal component analysis summarized morphometry of 10 variables (beak, wing, tarsus length, etc.). The overall body size (PC1) and the length of flight feathers regarding body size (PC3) showed significant positive relationships with plumage redness. Plumage redness was barely correlated with bill shape measures, suggesting no constraint in acquiring carotenoids from pine cones. However, large body sizes or proportionally long flying feathers could help carotenoid acquisition via social competition or increased foraging ranges. Proportionally longer flight feathers might also be associated with a specific cell respiration profile that would simultaneously favor flying capacities and enzymatic transformations needed for ketocarotenoid synthesis. Such a phenotypic profile would agree with the hypothesis of ketocarotenoid-based colors acting as individual quality index signals.

Effects of Non-Native Shrubs on Caterpillars and Shrubland-Dependent Passerines within Three Transmission Line Rights-of-Way in Southeastern New Hampshire

Caterpillars (Lepidoptera, Hymenoptera: Symphyta) are a critical food that breeding passerines use for raising young and serve as an important source of carotenoids that support nestling growth and feather pigmentation. When non-native, invasive plants grow abundant enough to reduce native plant diversity they can cause a reduction in caterpillar abundance. The goal of this study was to determine whether a reduction in caterpillars caused by non-native, invasive shrubs affects nestling health and/or the reproductive success of a shrubland-dependent passerine, Geothlypis trichas trichas (Common Yellowthroat). I first quantified caterpillar abundance on the dominant native and nonnative shrubs comprising Common Yellowthroat territories in shrubby transmission-line rights-of-way and estimated total caterpillar abundance in each territory based on the shrub species composition. I then determined if differences in shrub species composition and caterpillar abundance affected: (1) adult Common Yellowthroat reproductive success, (2) nestling Common Yellowthroat diet composition, or (3) nestling Common Yellowthroat growth rate, plasma carotenoids, and carotenoid-based plumage color. Nonnative, invasive shrubs did not reduce caterpillar abundance until they comprised ≥55% of the shrub volume and significantly reduced the diversity and abundance of native shrubs in bird territories. Differences in caterpillar abundance among sites did not result in differences in Common Yellowthroat productivity, but where non-native shrubs comprised ≥55% of the shrub volume, adults increased their frequency of feeding visits to nestlings and fed nestlings the greatest proportion of non-caterpillar prey. Daily surveys of shrubland bird presence/absence at study sites indicated that most shrubland-dependent bird species were absent where non-native shrubs comprised ≥55% of the shrub cover; such conditions likely equated to habitat loss for bird species less able than Common Yellowthroats to adapt to low shrub diversity and low caterpillar abundance.

Plumage colour saturation predicts long-term, cross-seasonal social dominance in a mutually ornamented bird

Dominance hierarchies are known to reduce agonistic interactions between individuals and may be influenced by differences in phenotypes such as body size, personality or cognition. Colour ornamentation, especially with melanin-based colours, can also be a badge of status signalling social dominance, but status signalling with carotenoid-based colours is little understood. Here we studied whether different phenotypes, including individual differences in carotenoid-based colour ornamentation, body size, personality-, cognition- and stress-related traits, predict social dominance in the highly social and mutually ornamented common waxbill, Estrilda astrild . We monitored aggressive interactions of waxbills living in seminatural conditions, in a large mesocosm, using data from a radiofrequency identification system in the nonbreeding and breeding seasons over 2 consecutive years. We found the position of individual waxbills in the dominance hierarchy repeatable across seasons and years. Furthermore, the steepness of the dominance hierarchy was greater in the breeding seasons, perhaps because of increased competition over mates or breeding resources. Contrary to previous work with waxbills in birdcages, here body size did not predict social dominance, perhaps because in our more natural setting aggressive encounters were mostly airborne, where large body size may not confer a fighting advantage and may even reduce manoeuvrability. Sex, reactive-to-proactive personality differences and traits related to cognition or stress did not predict social dominance either. Finally, red colour saturation, but not the size of the red patch, strongly predicted social dominance and may thus act as a badge of status. This is one of the few studies testing whether size or saturation of carotenoid-based plumage coloration indicates social dominance and suggests that saturation may more often signal dominance. • Waxbills have mildly steep but stable social hierarchies across years and seasons. • Saturation of carotenoid-based red plumage colour indicates social dominance. • Size of red plumage areas does not correlate with social dominance. • Sex, personality, cognition and size were not related to social dominance.

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

Abstract Historically, studies of condition-dependent signals in animals have been male-centric, but recent work suggests that female ornaments can also communicate individual quality (e.g., disease state, fecundity). There also has been a surge of interest in how urbanization alters signaling traits, but we know little about if and how cities affect signal expression in female animals. We measured carotenoid-based plumage coloration and coccidian (Isospora spp.) parasite burden in desert and city populations of house finches Haemorhous mexicanus to examine links between urbanization, health state, and feather pigmentation in males and females. In earlier work, we showed that male house finches are less colorful and more parasitized in the city, and we again detected such patterns in this study for males; however, urban females were less colorful, but not more parasitized, than rural females. Moreover, contrary to rural populations, we found that urban birds (regardless of sex) with larger patches of carotenoid coloration were also more heavily infected with coccidia. These results show that urban environments can disrupt condition-dependent color expression and highlight the need for more studies on how cities affect disease and signaling traits in both male and female animals.

Expression of and choice for condition-dependent carotenoid-based color in an urbanizing context

Urban environments create a unique suite of conditions, leading to changes in animal behavior, morphology, phenology, and physiology. Condition-dependent traits such as the carotenoid-based coloration offer a unique opportunity to assess the impacts of urbanization on organisms because they reflect the nutrition, health, or other resource-based attributes of their bearers and they play an essential role in intra and intersex interactions. To determine if and how the carotenoid-based coloration of male house finches (Haemorhous mexicanus) varies along a gradient of urbanization, we quantified the plumage coloration of more than 1000 individuals in urban, suburban, rural, and desert habitats over the course of 17 months. We also examined for the first time the preference of females for male plumage coloration across the urban-rural gradient, to test if and how female preferences varied relative to the plumage coloration displayed by males in their local population. We found that carotenoid-based coloration decreased along the gradient of urbanization, suggesting that the enzyme-driven metabolic conversion of dietary carotenoids into red carotenoids used to color plumage is sensitive to urban stressors. The stronger negative effect of urbanization on carotenoid-based plumage coloration during breeding than during molt and winter suggests that urbanization affects color fading rate, maybe through modifications of feather-degrading bacterial load. Finally, we have shown that urbanization influences female mate-choice behavior, suggesting that female color preferences may track the variation in male coloration across the gradient of urbanization.

Plumage coloration and social context influence male investment in song.

Animals use multiple signals to attract mates, including elaborate song, brightly coloured ornaments and physical displays. Female birds often prefer both elaborate male song and intense carotenoid-based plumage coloration. This could lead less visually ornamented males to increase song production to maximize their attractiveness to females. We tested this possibility in the highly social and non-territorial house finch (Haemorhous mexicanus), in which females discriminate among males based on both song and on the intensity of red carotenoid-based plumage coloration. We manipulated male plumage coloration through carotenoid supplementation during moult, so that males were either red or yellow. Males were then housed under three social environments: (i) all red birds, (ii) all yellow birds or (iii) a mixture of red/yellow birds. We recorded song after presentation of a female. Red males produced more song than yellow males. But when yellow males were housed with red conspecifics, they produced more song relative to yellow males housed with equally unattractive yellow males. This study provides novel evidence that a male's plumage coloration and the plumage colour of his social competitors influence investment in song.

Effects of an early-life paraquat exposure on adult resistance to oxidative stress, plumage colour and sperm performance in a wild bird.

Early-life stressful conditions can shape individual phenotypes and ultimately influence fitness. Oxidative stress is a pervasive threat that affects many fitness-related traits and can modulate life-history trade-offs. Yet, the extent to which exposure to oxidative stress during early life can have long-lasting effects on key fitness-related traits remains to be elucidated, particularly in natural populations of vertebrates. Using a wild population of great tits Parus major, we experimentally dosed 11-day-old birds with paraquat, a pro-oxidant molecule, aiming at increasing oxidative stress. One year later, we recaptured 39 of them as adult recruiting breeders and quantified effects of the paraquat exposure on their resistance to oxidative stress, carotenoid-based plumage coloration and male sperm performance. Despite the absence of a short-term effect of paraquat on oxidative stress measured two days later, the pre-fledging exposure to paraquat induced a reduction in individual oxidative damage measured at adulthood. Paraquat-dosed individuals also had brighter plumage, but no effect was observed on male sperm performance. For the first time in a natural population of vertebrates, we experimentally show that an early-life acute exposure to a pro-oxidant has long-lasting effects on individual resistance to oxidative stress at adulthood. Our results are in line with the environmental matching and the hormesis hypotheses but may also reflect selective disappearance of individuals with lower resistance to oxidative stress.

Biogeographical variation of plumage coloration in the sexually dichromatic Hawai‘i ‘Amakihi (Chlorodrepanis virens)

Plumage coloration in birds can be of major importance to mate selection, social signaling, or predator avoidance. Variations in plumage coloration related to sex, age class, or seasons have been widely studied, but the effect of other factors such as climate is less known. In this study, we examine how carotenoid-based plumage coloration and sexual dichromatism of the Hawai‘i ‘Amakihi (Chlorodrepanis virens) varies with rainfall and temperature on Hawai‘i Island. We also examined whether Hawai‘i ‘Amakihi plumage coloration patterns follow Gloger’s rule, which states that animals in wetter climates have darker coloration. Hawai‘i ‘Amakihi were mist-netted and banded at 12 sites representing six major climatic zones on Hawai‘i Island. Feather samples were collected from two body regions: the breast and rump. Using spectrophotometry, we recorded coloration using measures of hue, saturation, and brightness. We conducted sex determination by polymerase chain reaction to confirm the sex of birds sampled. We found that the plumage coloration of Hawai‘i ‘Amakihi varied with both temperature and rainfall. ‘Amakihi plumage’s brightness showed a quadratic relationship with rainfall, contrary to Gloger’s rule, and decreased with temperature. Saturation depended on the interaction between temperature and rainfall. Increases in rainfall also increased saturation in warm areas, while they reduced saturation when the temperature was low. Finally, we found chromatic differences among sexes, but sexual dichromatism was not affected by the climatic conditions. This study provides evidence that rainfall and temperature play an important role in determining the plumage traits of Hawai‘i ‘Amakihi.

Are long-term widespread avian body size changes related to food availability? A test using contemporaneous changes in carotenoid-based color.

Recent changes in global climate have been linked with changes in animal body size. While declines in body size are commonly explained as an adaptive thermoregulatory response to climate warming, many species do not decline in size, and alternative explanations for size change exist. One possibility is that temporal changes in animal body size are driven by changes in environmental productivity and food availability. This hypothesis is difficult to test due to the lack of suitable estimates that go back in time. Here, we use an alternative, indirect, approach and assess whether continent‐wide changes over the previous 100 years in body size in 15 species of Australian birds are associated with changes in their yellow carotenoid‐based plumage coloration. This type of coloration is strongly affected by food availability because birds cannot synthesize carotenoids and need to ingest them, and because color expression depends on general body condition. We found significant continent‐wide intraspecific temporal changes in body size (wing length) and yellow carotenoid‐based color (plumage reflectance) for half the species. Direction and magnitude of changes were highly variable among species. Meta‐analysis indicated that neither body size nor yellow plumage color showed a consistent temporal trend and that changes in color were not correlated with changes in size over the past 100 years. We conclude that our data provide no evidence that broad‐scale variation in food availability is a general explanation for continent‐wide changes in body size in this group of species. The interspecific variability in temporal changes in size as well as color suggests that it might be unlikely that a single factor drives these changes, and more detailed studies of museum specimens and long‐term field studies are required to disentangle the processes involved.

Conditions on the Mexican moulting grounds influence feather colour and carotenoids in Bullock's orioles (Icterus bullockii)

Carotenoid‐based plumage coloration plays a critical role for both inter‐ and intrasexual communication. Habitat and diet during molt can have important consequences for the development of the ornamental signals used in these contexts. When molt occurs away from the breeding grounds (e.g., pre‐alternate molt on the wintering grounds, or stopover molt), discerning the influence of habitat and diet can be particularly important, as these effects may result in important carryover effects that influence territory acquisition or mate choice in subsequent seasons. Several species of songbirds in western North America, including the Bullock's oriole (Icterus bullockii), migrate from the breeding grounds to undergo a complete prebasic (post‐breeding) molt at a stopover site in the region affected by the Mexican monsoon climate pattern. This strategy appears to have evolved several times independently in response to the harsh, food‐limited late‐summer conditions in the arid West, which contrast strongly with the high productivity driven by heavy rains that is characteristic of the Mexican monsoon region. Within this region, individuals may be able to optimize plumage coloration by molting in favourable areas characterized by high resource abundance. We used stable isotope analysis (δ13C, δ15N) to ask whether the diet and molt habitat/location of Bullock's orioles influenced their expression of carotenoid‐based plumage coloration as well as plumage carotenoid content and composition. Bullock's orioles with lower feather δ15N values acquired more colorful plumage (orange‐shifted hue) but had feathers with lower total carotenoid concentration, lower zeaxanthin concentration, and marginally lower canthaxanthin and lutein concentration. Examining factors occurring throughout the annual cycle are critical for understanding evolutionary and ecological processes. Here, we demonstrate that conditions experienced during a stopover molt, occurring hundreds to thousands of kilometers from the breeding grounds, influence the production of ornamental plumage coloration, which may carryover to influence inter‐ and intrasexual signaling in subsequent seasons.

Effects of diet on plumage coloration and carotenoid deposition in red and yellow domestic canaries (Serinus canaria)

ABSTRACT Atlantic Canaries (Serinus canaria) are the most commonly kept caged bird with extensive carotenoid-based plumage coloration. Domestic strains of canaries have been bred for a variety of colors and patterns, making them a valuable model for studies of the genetic bases for feather pigmentation. However, no detailed account has been published on feather pigments of the various strains of this species, particularly in relation to dietary pigments available during molt. Moreover, in the twentieth century, aviculturists created a red canary by crossing Atlantic Canaries with Red Siskins (Carduelis cucullata). This “red-factor” canary is reputed to metabolically transform yellow dietary pigments into red ketocarotenoids, but such metabolic capacity has yet to be documented in controlled experiments. We fed molting yellow and red-factor canaries seed diets supplemented with either β-carotene, lutein/zeaxanthin, or β-cryptoxanthin/β-carotene and measured the coloration and carotenoid content of newly gr...

How to reduce the costs of ornaments without reducing their effectiveness? An example of a mechanism from carotenoid-based plumage.

Carotenoid-based ornaments are often considered to be honest indicators of individual quality assessed by potential mates. However, males can use a variety of strategies that minimize the amount of costly carotenoids used while retaining the effectiveness of color signaling. Birds could do this by altering pigment intake, metabolism, or its presentation to a potential signal receiver. Here, we propose a new mechanism of lowering the costs of carotenoid displays in birds: differential allocation of pigments within single feathers. We studied the coloration of the yellow terminal tail bands of rectrices of male Bohemian waxwings. Using reflectance spectrometry, we show that the two central rectrices are most intensively colored compared to other rectrices. More detailed analyses reveal that these differences result from feather-specific patterns of rectrices coloration. The outer feather vanes of the outermost rectrices are more intensively colored compared to the inner vanes. However, the central rectrices have equally colored vanes that are, on average, more intensively pigmented than the outermost rectrices. When the waxwing tail is folded, the outermost rectrices are covered by other feathers, except for the narrow, outer vane. Central rectrices, however, form the outermost layers which are not obscured by other tail feathers. Thus, the feather vanes that are the most visible to potential viewers are also the most pigmented. These results support the occurrence of a previously overlooked mechanism to reduce the costs of carotenoid-based ornaments: precise pigment distribution to maximize efficiency of signals within single feathers.Significance statementMales of many bird species use bright carotenoid-based plumage coloration to attract females. These traits are physiologically expensive such that only individuals in prime condition can develop the most vivid colors. Males often “cheat” to obtain attractive appearances at lower costs. We showed that this goal could be achieved by differential deposition of pigments into the most conspicuous feather regions. Bohemian waxwing males have yellow tips on their rectrices of which the outer vanes are more brightly colored compared to the inner vanes. These inner feather vanes are usually covered by other feathers and are, thus, less visible to conspecifics. The only exception is the pair of central rectrices that are fully exposed, and both feather vanes are equally colored. In this species, males minimize the use of costly carotenoid pigments while maintaining elaborate ornamentation of plumage regions that are most visible to potential mates.

Evidence for multiple functions in a sexually selected ornament

In gregarious animals, social interactions frequently take the form of dominance hierarchies that maintain stable relationships between individuals, and settle disputes without extra costs. Traits that function as signals of status can play an important role in mediating interactions among individuals, both in social and in sexual contexts. Carotenoid ornaments are more generally assumed to be sexually selected and not so relevant to general social contexts. However, it is possible for them to function in social contexts if they signal socially relevant aspects. Here we experimentally analysed social dominance and resource control in male groups of a gregarious species, the European serin, Serinus serinus, in relation to a sexual ornament. We tested whether yellow carotenoid-based plumage coloration, age, body size and testosterone were predictors of social dominance over a nonsexual resource (i.e. feeding context). We showed that dominance hierarchies were steep and were related to testosterone levels and ornamental coloration, particularly the male yellow carotenoid-based crown patch. Our results suggest that carotenoid-based colour and testosterone levels can be reliable predictors of social status in agonistic encounters in groups of male serins. Moreover, together with previous work on the sexual function of male coloration, this study provides evidence that male serin yellow coloration has a dual function in both sexual and nonsexual contexts. These results raise the possibility that this ornament may have evolved and be maintained via social selection over social competition/cooperation for reproductive opportunities and ecological resources.

Is oxidative status influenced by dietary carotenoid and physical activity after moult in the great tit (Parus major)?

In the context of sexual and natural selection, an allocation trade-off for carotenoid pigments may exist because of their obligate dietary origin and their role both in the antioxidant and immune systems and in the production of coloured signals in various taxa, particularly birds. When birds have expended large amounts of carotenoids to feather growth such as after autumn moult, bird health and oxidative status might be more constrained. We tested this hypothesis in a bird species with carotenoid-based plumage colour, by manipulating dietary carotenoids and physical activity, which can decrease antioxidant capacity and increase reactive oxygen metabolite (ROM) concentration. Great tits were captured after moult and kept in aviaries, under three treatments: physical handicap and dietary supplementation with carotenoids, physical handicap and control diet, and no handicap and control diet. We measured plasma composition (antioxidant capacity, ROM concentration, and vitamin A, vitamin E and total carotenoid concentrations), immune system activation (blood sedimentation) and stress response (heterophil/lymphocyte ratio) and predicted that handicap treatment should influence these negatively and carotenoid supplementation positively. Coloration of yellow feathers was also measured. Carotenoid supplementation increased total plasma carotenoid concentration, decreased feather carotenoid chroma and marginally increased ROM concentration. Handicap increased blood sedimentation only in males but had no clear influence on oxidative stress, which contradicted previous studies. Further studies are needed to investigate how physical activity and carotenoid availability might interact and influence oxidative stress outside the moult period, and their combined potential influence on attractiveness and reproductive investment later during the breeding season.

Habitat structure and prey composition generate contrasting effects on carotenoid-based coloration of great titParus majornestlings

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.