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.     

Energetic constraints on expression of carotenoid-based plumage coloration

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

Air pollution fades the plumage of the Great Tit

1. Great Tits ( Parus major ) derive the carotenoid pigments for their yellow plumage via the prey items in their diet. Air pollution is known to affect the abundance of many forest insects, e.g. green caterpillars, which are an important source of food and pigments for tits. This study investigates whether air pollutants indirectly affected the intensity of the yellow colour in P. major plumage via the reduced access to carotenoid sources.
2. The intensity of the yellow colour in the plumage of P. major nestlings was scored and the relative abundance of green herbivorous larvae in territories around a polluting copper smelter in SW Finland was simultaneously measured.
3. Both the intensity of yellow colour in nestling plumage and caterpillar abundance increased with increasing distance from the pollution source. The colour intensity correlated significantly with the density of green herbivorous larvae in a territory.
4. Parus major nestlings were significantly heavier at distant sites than close to the pollution source which suggests that the future survival probability of pale nestlings may be lowered.
5. Young birds, after their first moult, were studied for the relationships between condition, size and plumage colour by the means of ptilochronology. The plumage colour intensity did not correlate with the size corrected width of the growth bars in fifth rectrix (condition at moult) but was correlated positively with the length of the rectrix (size).
6. The implications of colour change for survival and mate choice are discussed.     

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

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

人工巢箱条件下的大山雀巢捕食

于2004—2008年在次生阔叶林中,采用悬挂巢箱的方法对大山雀的巢捕食作了研究。结果表明:不同年龄巢箱的被捕食率显著不同,新巢箱被捕食率最低,第2年被捕食率最高,第3年下降很大,第4年又略有上升。被捕食巢的窝卵数极显著的低于未被捕食巢的窝卵数。影响巢捕食的主要生境因子为巢箱高度和巢上盖度,其次为灌木的密度和高度。     

The carotenoid-based plumage coloration of adult Blue Tits Cyanistes caeruleus correlates with the health status of their brood

The Blue Tit Cyanistes caeruleus is a passerine bird in which both sexes provide substantial care to the offspring and display conspicuous carotenoid-based plumage coloration. It has been shown that carotenoid-based coloration in birds reflects both individual quality and foraging ability. Because the body condition of nestlings usually depends on the capacity of their parents to feed them, I predicted that, independent of sex, those individuals with the most exaggerated carotenoid-based plumage coloration should raise offspring in better health. I found that although, in my study population, the smallest females were paired with the largest males, the brightest females were paired with the brightest and most intensely coloured males. I used body condition and T-cell-mediated immune response of nestlings as measures of their health status. Generalized mixed models showed that brighter and more-yellow adults reared offspring in better than average condition and immune response. Older males and smaller females were also able to raise offspring with better immune response. All these results suggest that the carotenoid-based plumage coloration of parents is somehow linked with individual quality, as it presents a significant and positive correlation with the health status of their growing chicks. Thus, the brightest and more intensely coloured individuals raised the healthiest offspring.     

Proximate mechanisms of variation in the carotenoid-based plumage coloration of nestling great tits (Parus major L.)

Many vertebrates use carotenoid-based signals in social or sexual interactions. Honest signalling via carotenoids implies some limitation of carotenoid-based colour expression among phenotypes in the wild, and at least five limiting proximate mechanisms have been hypothesized. Limitation may arise by carotenoid-availability, genetic constraints, body condition, parasites, or detrimental effects of carotenoids. An understanding of the relative importance of the five mechanisms is relevant in the context of natural and sexual selection acting on signal evolution. In an experimental field study with carotenoid supplementation, simultaneous cross-fostering, manipulation of brood size and ectoparasite load, we investigated the relative importance of these mechanisms for the variation in carotenoid-based coloration of nestling great tits (Parus major). Carotenoid-based plumage coloration was significantly related to genetic origin of nestlings, and was enhanced both in carotenoid-supplemented nestlings, and nestlings raised in reduced broods. We found a tendency for ectoparasite-induced limitation of colour expression and no evidence for detrimental effects of carotenoids on growth pattern, mortality and recruitment of nestlings to the local breeding population. Thus, three of the five proposed mechanisms can generate individual variation in the expression of carotenoid-based plumage coloration in the wild and thus could maintain honesty in a trait potentially used for signalling of individual quality.     

人工巢箱中大山雀和褐头山雀的繁殖比较

2009年3-7月,在北京小龙门森林公园悬挂人工巢箱,对大山雀(Parus major)和褐头山雀(P.montanus;)的繁殖进行比较研究.野外共悬挂100个巢箱,其中19巢(19.0%)被大山雀、11巢(11.0%)被褐头山雀进驻,总利用率为30.0%.同域繁殖的大山雀和褐头山雀在窝卵数、孵化期上有极显著差异(P＜0.01),其中大山雀的窝卵数(8.21枚±0.25枚,n=19)极显著大于褐头山雀(6.18枚±0.23枚,n=11),而褐头山雀的孵化期(14.22d±0.44d,n=10)极显著长于大山雀(13.17 d±O.83 d,n=12).两者在卵重、卵大小、出雏数、雏鸟出飞数上无显著差异(P＞0.05).大山雀的繁殖成功率(27.0%)和营巢成效(63.2%)均显著低于褐头山雀(54.5%和100%)(P＜0.01).表明大山雀可能采取高产卵数、低存活率的繁殖对策,而褐头山雀的繁殖策略则可能为低产卵数、高存活率.     

Measurement of plumage badges: an evaluation of methods used in the Great Tit Parus major

There is an increasing interest in the variation in plumage coloration and a number of theories have been proposed to explain its evolution (Butcher & Rohwer 1989, Savalli 1995). Birds haye been considered to signal through plumage their dominance status, ability to evade predators, parasite resistance or for example their parental care abilities (reviewed in Butcher & Rohwer 1989, Anderson 1994, Savalli 1995, Johnstone 1997). However, the measuremcnt of plumage badges and its accuracy have received little attention (Savalli 1995). In the case of the Great Tit Parus major, for instance, several studies have analysed the importance of the breast stripe size as an indicator of dominance status in winter flocks (Jarvi & Bakken 1984, Piiysa 1988, Lemel 1989, Wilson 1992) and its importance in mate choice (Norris 1990, 1993), but the badgc has been estimated using very different indices, even in studies performed by the same authors (Norris 1990, 1993). We compare the performance of differvnt indices in estimating Great Tit stripe size, by comparing them with badge surface area obtained from image analysis of digital photographs. We propose that digital photography provides a reliable and inexpensive method to measure plumage badges in birds and we demonstrate its performance in the Great Tit.     

Testing the shared-pathway hypothesis in the carotenoid-based coloration of red crossbills

The mechanisms involved in the production of red carotenoid-based ornaments of vertebrates are still poorly understood. These colorations often depend on enzymatic transformations (ketolation) of dietary yellow carotenoids, which could occur in the inner mitochondrial membrane (IMM). Thus, carotenoid ketolation and cell respiration could share biochemical pathways, favoring the evolution of ketocarotenoid-based ornaments as reliable indices of individual quality under sexual selection. Captive male red crossbills (Loxia curvirostra Linnaeus) were exposed to redox-active compounds designed to penetrate and act in the IMM: an ubiquinone (mitoQ) or a superoxide dismutase mimetic (mitoTEMPO). MitoQ can act as an antioxidant but also distort the IMM structure, increasing mitochondrial free radical production. MitoQ decreased yellow carotenoids and tocopherol levels in blood, perhaps by being consumed as antioxidants. Contrarily, mitoTEMPO-treated birds rose circulating levels of the second most abundant ketocarotenoid in crossbills (i.e., canthaxanthin). It also increased feather total red ketocarotenoid concentration and redness, but only among those birds exhibiting a redder plumage at the start of the study, that is, supposedly high-quality individuals. The fact that mitoTEMPO effects depended on original plumage color suggests that the red-ketocarotenoid-based ornaments indicate individual quality as mitochondrial function efficiency. The findings would thus support the shared pathway hypothesis.     

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

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

The relationship between plumage coloration and aggression in female tree swallows

Intrasexual competition is an important selective force that can favor the evolution of honest signals of fighting ability. Research has focused predominantly on male birds, but many female birds also possess plumage ornaments that could mediate the outcome of competitive interactions. We examined the relationship between blue and white structural coloration and aggression in female tree swallows Tachycineta bicolor. Tree swallows are secondary cavity nesters and females show delayed plumage maturation which may be related to intense competition for nest sites. We compared plumage reflectance of second‐year (SY) and after‐second year (ASY) females, and within‐individual changes in plumage reflectance of ASY females in two successive years. We assessed aggression by placing a caged SY female 2 m from the nest box of an ASY female and quantified the ASY female's response in relation to her own plumage coloration. We found substantial differences in plumage reflectance of SY and ASY females, but found that ASY females became greener with duller white coloration in the second year. This may be due to poor weather that made reproduction particularly costly in the first year of the study and suggests coloration is influenced by condition. We found no relationship between dorsal coloration and aggressiveness. Rather, brighter white females spent more time on their nest box and within 2 m of the intruder than females with dull breasts. Our findings suggest that brighter white ASY females may perceive a SY female as less of a threat or that there may be a trade‐off associated with aggression and parental care that is related to white brightness.     

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

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

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

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

Selection and inheritance of sexually dimorphic juvenile plumage coloration

Sexually dimorphic plumage coloration is widespread in birds and is generally thought to be a result of sexual selection for more ornamented males. Although many studies find an association between coloration and fitness related traits, few of these simultaneously examine selection and inheritance. Theory predicts that sex‐linked genetic variation can facilitate the evolution of dimorphism, and some empirical work supports this, but we still know very little about the extent of sex linkage of sexually dimorphic traits. We used a longitudinal study on juvenile Florida scrub‐jays (Aphelocoma coerulescens) to estimate strength of selection and autosomal and Z‐linked heritability of mean brightness, UV chroma, and hue. Although plumage coloration signals dominance in juveniles, there was no indication that plumage coloration was related to whether or not an individual bred or its lifetime reproductive success. While mean brightness and UV chroma are moderately heritable, hue is not. There was no evidence for sex‐linked inheritance of any trait with most of the variation explained by maternal effects. The genetic correlation between the sexes was high and not significantly different from unity. These results indicate that evolution of sexual dimorphism in this species is constrained by low sex‐linked heritability and high intersexual genetic correlation.     

The evolution of bill coloration and plumage dimorphism supports the transference hypothesis in dabbling ducks

Bright coloration in male birds is typically thought to be drivenby sexual selection (female choice or male-male competition).Bird species often vary in the intensity of bright coloration,but few studies have addressed this cross-species variation.Potentially this variation could result from either variationin female preferences or in the relative costs of male traits. Speciesof dabbling ducks vary in the presence of bright male plumageand bill coloration. I tested the transference hypothesis forornament evolution in dabbling ducks using a phylogenetic studyof character evolution. The transference hypothesis makes threepredictions: (1) a costly male ornamental trait is the ancestralcondition, (2) a less costly male ornamental trait is the derivedcondition, and (3) gains in the less costly trait are associatedwith losses or absence of the more costly male trait. All threeof these predictions were satisfied in this study of the evolutionof plumage dimorphism and bright bill coloration in the dabblingducks, given that bright plumage coloration is more costly thanbright bill coloration.     

The colour of fitness: plumage coloration and lifetime reproductive success in the tawny owl

We studied variation in plumage colour and life history in a population of tawny owls (Strix aluco) in southern Finland, using 26 years of data on individually marked male and female owls. Colour was scored on a semi-continuous scale from pale grey to reddish brown. Colour scoring was repeatable and showed a bimodal distribution (grey and brown morph) in both sexes. During the study period, colour composition was stable in the study population in both sexes. The sexes did not mate assortatively with respect to their colour. Colour was a highly heritable trait and was under selection. Grey-coloured male and female owls had a higher lifetime production of fledglings, and grey-coloured male (but not female) owls produced more recruits during their lifetime than brown individuals. Selection on colour was mediated through viability selection and not through fecundity selection. Our results reveal remarkably strong selection on a genetically determined phenotypic trait.