Plumage Ornaments Research Articles

Major breeding plumage color differences of male ruffs (Philomachus pugnax) are not associated with coding sequence variation in the MC1R gene.

Sequence variation in the melanocortin-1 receptor (MC1R) gene explains color morph variation in several species of birds and mammals. Ruffs (Philomachus pugnax) exhibit major dark/light color differences in melanin-based male breeding plumage which is closely associated with alternative reproductive behavior. A previous study identified a microsatellite marker (Ppu020) near the MC1R locus associated with the presence/absence of ornamental plumage. We investigated whether coding sequence variation in the MC1R gene explains major dark/light plumage color variation and/or the presence/absence of ornamental plumage in ruffs. Among 821bp of the MC1R coding region from 44 male ruffs we found 3 single nucleotide polymorphisms, representing 1 nonsynonymous and 2 synonymous amino acid substitutions. None were associated with major dark/light color differences or the presence/absence of ornamental plumage. At all amino acid sites known to be functionally important in other avian species with dark/light plumage color variation, ruffs were either monomorphic or the shared polymorphism did not coincide with color morph. Neither ornamental plumage color differences nor the presence/absence of ornamental plumage in ruffs are likely to be caused entirely by amino acid variation within the coding regions of the MC1R locus. Regulatory elements and structural variation at other loci may be involved in melanin expression and contribute to the extreme plumage polymorphism observed in this species.

Plumage ornaments and reproductive investment in relation to oxidative status in the Iberian Pied Flycatcher (Ficedulahypoleuca iberiae)

A key aspect in the study of plumage traits with a potential role in communication is the cost associated with trait production and maintenance, expressed in terms of oxidative stress. In the Iberian Pied Flycatcher (Ficedula hypoleuca iberiae (Witherby, 1928)), males and some females exhibit a white forehead patch and both sexes present conspicuous white patches on the wings. We examined associations between these plumage ornaments and their ability to cope with oxidative stress. Furthermore, we explored oxidative costs of reproductive investment. Total antioxidant status (TAS) in plasma and glutathione (tGSH) levels in red blood cells, as well as a marker of oxidative damage in plasma lipids (malondialdehyde (MDA)), were assessed simultaneously for the first time in studies of avian reproduction. We found negative associations between antioxidants and ornaments in incubating females, although this relationship was positive while feeding nestlings. For males, MDA levels were negatively associated with ornaments, while TAS showed a positive relation. Female MDA showed a positive correlation with intensity of incubation attendance, while this relation was negative for tGSH levels. These results indicate that multiple achromatic plumage ornaments signal the individual capacity to cope with costs related to oxidative stress. Moreover, this study highlights the critical role of incubation for avian life histories.

Steller sex: infidelity and sexual selection in a social Corvid (Cyanocitta stelleri).

Genetic analysis of avian mating systems has revealed that more than 70% of monogamous species show incidence of offspring parentage that does not match the social partner. Extra-pair parentage (EPP) has been linked to a variety of factors, including size and symmetry of ornamental traits, coloration, resource availability, and local conspecific density. We examined how ornamental plumage traits of individual Steller's jays (Cyanocitta stelleri) and territory characteristics influence genetic fidelity of socially monogamous pairs. We used seven highly polymorphic microsatellite markers to assign paternity to 79 offspring, and identified 12 (15.2%) as extra-pair young (EPY). Steller's jays with extra-pair young had significantly lower values of feather brightness and hue, indicating more ultraviolet-blue shifted coloration, and nested in closer proximity to the forest edge than Steller's jays with no detected EPY. Body size, crest height, asymmetry of ornamental crest stripes, as well as vegetative composition of territories and their proximity to supplemental feeders appeared to have little relationship to EPP. These results indicate that extra-pair parentage plays a role in the evolution of secondary sexual characteristics in both sexes, and suggest local density and availability of resources may influence Steller's jay mating dynamics.

Plumage ornaments in male Northern Lapwings Vanellus vanellus may not be a reliable indicator of their mating success

Conspicuous morphological traits that signal aspects of physical state of the owner may be used as cues in mate choice, leading to sexual selection of ornamental traits. In birds, colour patterns as well as structural attributes may constitute ornaments affecting mate choice. In the breeding season, Northern Lapwing Vanellus vanellus males possess long crests, contrast-rich black and white facial markings, a black breast, and a shining greenish dorsal side. In non-breeding and juvenile plumages, and in females, crests are shorter, facial patterns have less clear contrasts, and breast and dorsal side possess varying amounts of light fringes. Depending onmoult energetics in latewinter/early spring, when males acquire their breeding plumage, the state of these characters could potentially signal male condition to females prospecting for mates. We examined if these traits correlated with male mating success in this polygynous species, predicting males with the most expressed ornamental characters (a) tomate earlier and (b) to gain more mates.Trait states were ranked for 31 males of which breeding phenology and number of mates were known. No correlationwas found between trait expression and date of first egg laid on the territory and number of mates did not correlate with trait expression. Thus, the study did not confirm that females use crest length, breast blackness and face contrasts or lack of dorsal fringes as cues in their mate choice. This could be because the elaborate traits do not have an ornamental function important for femalemate choice, but experimental studies would be needed to rigorously test this hypothesis.

Contrasting patterns of selection on the size and coloration of a female plumage ornament in common yellowthroats

Females often possess ornaments that appear smaller and duller than homologous traits in males. These ornaments may arise as nonfunctional by-products of sexual selection in males and cause negative viability or fecundity selection in females in proportion to the cost of their production and maintenance. Alternatively, female ornaments may function as signals of quality that are maintained by sexual or social selection. In a 4-year study of 83 female common yellowthroats (Geothlypis trichas) and their 222 young, we found strong viability and fecundity selection on the yellow bib, a carotenoid-based plumage ornament that is a target of sexual selection in males. Females with larger bibs were older, larger and more fecund than females with smaller bibs. However, bib size positively covaried with bib total brightness and carotenoid chroma, aspects of bib coloration that were under negative viability and fecundity selection. Females with more colourful bibs laid fewer eggs in their first clutch, were more likely to suffer total brood loss due to predation and were less likely to return to the study area. Selection against bib coloration limits the value of bib size as a quality indicator in females and may constrain the elaboration of bib attributes in males.

Contemporary divergence of island bird plumage

Although the diversity in avian plumage coloration is striking, there is little known about the rate with which colour diverges. Eastern bluebirds Sialia sialis bermudensis on the island of Bermuda are considered endemic based upon differences in coloration from the mainland, but recent molecular evidence suggests they established on the island only 400 yr ago. We explored sexual dichromatism and colour divergence in this isolated population, thus providing one of the few quantitative accounts of contemporary plumage change. Contrary to expectations that sexual dichromatism would decrease in this sedentary island population, we found that males and females have increased plumage ornamentation in a coordinated fashion that acts to preserve sexual dichromatism, while plumage colour is also altered to become brighter and bluer. These differences were in place at least 100 yr ago based upon a separate analysis of museum specimens. Our results provide insight into the divergence of plumage colour in an incipient species, and we show the remarkable extent to which plumage colour can change over contemporary time frames.

The separate effects of egg size and parental quality on the development of ornamental plumage coloration

The fitness‐related consequences of egg size, independent of the influences of parental quality, are poorly understood in altricial birds. Not only can egg size and parental quality influence growth and survival, but each could influence the development of condition‐dependent plumage coloration in offspring. The Eastern Bluebird Sialia sialis is an altricial, multi‐brooded, cavity‐nesting passerine in which juveniles display dichromatic UV‐blue plumage. Previous research suggests that plumage coloration acts as a signal of individual quality among juvenile and adult Eastern Bluebirds. Here, we separate the effects of egg size and parental quality (defined by egg size laid) on nestling growth and plumage ornamentation by exchanging clutches of large eggs with clutches of small eggs. Nestlings were significantly larger immediately post‐hatching when hatched from a large egg, but to maintain a larger size, nestlings needed to have hatched from a large egg and to have been reared by high‐quality parents. Nestlings were brighter when reared by high‐quality parents and this relationship was strongest later in the breeding season. Nestlings exhibited greater UV chroma if hatched early in the season, but UV chroma was not significantly affected by egg size or parental quality. These findings demonstrate varying influences of both egg size and parental quality on offspring growth and plumage ornamentation but suggest that quality of post‐hatching investment is more influential than pre‐hatching investment.

Phylogenetics and diversification of tanagers (Passeriformes: Thraupidae), the largest radiation of Neotropical songbirds

Thraupidae is the second largest family of birds and represents about 4% of all avian species and 12% of the Neotropical avifauna. Species in this family display a wide range of plumage colors and patterns, foraging behaviors, vocalizations, ecotypes, and habitat preferences. The lack of a complete phylogeny for tanagers has hindered the study of this evolutionary diversity. Here, we present a comprehensive, species-level phylogeny for tanagers using six molecular markers. Our analyses identified 13 major clades of tanagers that we designate as subfamilies. In addition, two species are recognized as distinct branches on the tanager tree. Our topologies disagree in many places with previous estimates of relationships within tanagers, and many long-recognized genera are not monophyletic in our analyses. Our trees identify several cases of convergent evolution in plumage ornaments and bill morphology, and two cases of social mimicry. The phylogeny produced by this study provides a robust framework for studying macroevolutionary patterns and character evolution. We use our new phylogeny to study diversification processes, and find that tanagers show a background model of exponentially declining diversification rates. Thus, the evolution of tanagers began with an initial burst of diversification followed by a rate slowdown. In addition to this background model, two later, clade-specific rate shifts are supported, one increase for Darwin’s finches and another increase for some species of Sporophila. The rate of diversification within these two groups is exceptional, even when compared to the overall rapid rate of diversification found within tanagers. This study provides the first robust assessment of diversification rates for the Darwin’s finches in the context of the larger group within which they evolved.

Revisiting the condition‐dependence of melanin‐based plumage

Support against the condition‐dependence of melanin plumage signals has relied on data from species exhibiting both melanin‐ and carotenoid‐based plumage ornaments. As the mechanisms leading to variation in carotenoid‐ and melanin‐based plumage differ fundamentally, these systems may not be ideal to assess the condition‐dependence of melanin signals. Instead, we hypothesized that melanin‐plumage is more likely to signal condition in purely achromatic species. We performed a meta‐analysis reviewing evidence for condition‐dependent melanic plumage: we compared the net effect size for the relationship between melanin traits and condition in species that are achromatic versus species that also display a carotenoid‐based trait. Our results indicate that melanin plumage is condition‐dependent in species of both plumage types. Contrarily to our prediction, this finding suggests that melanin ornament condition‐dependence is not conditional on the context of other ornaments within a species. Instead, melanin ornaments should be viewed as potential condition‐dependent signals in all species.

The size of a melanin-based plumage ornament correlates with glucocorticoid receptor concentrations in the skin of that ornament

Glucocorticoid hormones such as corticosterone (CORT) play crucial roles in many physiological processes. CORT's actions are primarily mediated via binding to two receptors (glucocorticoid receptors (GRs) and mineralocorticoid receptors (MRs)) in different target tissues. CORT receptors can be independently regulated from circulating hormone titres, from tissue to tissue and even within different regions of the same tissue type. Increasing evidence has shown relationships between circulating CORT and melanin-based pigmentation in skin and feathers, yet to our knowledge, there have been no studies of CORT receptors in the skin of melanized ornaments. Male house sparrows (Passer domesticus) have a black, melanized bib, and evidence suggests that bib size is an important intraspecific signal. We examined the relationship between bib area and tissue sensitivity to CORT by quantifying GR and MR in bib skin and in adjacent paler-feathered belly skin (as a control tissue) at different life-history stages using radioligand binding assays. Males with larger bibs relative to their life-history stage had less GR in bib skin, but not belly skin, than males with smaller bibs. These results suggest a connection between the size of a melanin-based ornament and the underlying tissue's responsiveness to CORT.

Integrated plumage colour variation in relation to body condition, reproductive investment and laying date in the collared flycatcher

The possible integration of different sexual ornaments into a composite system, and especially the information content of such ornament complexes, is poorly investigated. Many bird species display complex plumage coloration, but whether this represents one integrated or several independent sexual traits can be unclear. Collared flycatchers (Ficedula albicollis) display melanised and depigmented plumage areas, and the spectral features (brightness and UV chroma) of these are correlated with each other across the plumage. In a 5-year dataset of male and female plumage reflectance, we examined some of the potential information content of integrated, plumage-level colour attributes by estimating their relationships to previous and current year body condition, laying date and clutch size. Females were in better condition the year before they became darker pigmented, and males in better current year condition were also darker pigmented. Female pigment-based brightness was positively, while male structurally based brightness was negatively related to current laying date. Finally, the overall UV chroma of white plumage areas in males was positively associated with current clutch size. Our results show that higher degree of pigmentation is related to better condition, while the structural colour component is associated with some aspects of reproductive investment. These results highlight the possibility that correlated aspects of a multiple plumage ornamentation system may reflect together some aspects of individual quality, thereby functioning as a composite signal.

The carotenoid‐based red cap of theMiddleSpottedWoodpeckerDendrocopos mediusreflects individual quality and territory size

Carotenoid‐based plumage ornaments have the potential to signal individual condition and health in many species of birds. However, very little is known about the function of red plumage in woodpeckers. We assessed whether the red cap displayed by both male and femaleMiddleSpottedWoodpeckers reflects individual quality, finding that the size of the cap is sex‐dependent, whereas the brightness of the cap correlates with the body condition of an individual. Furthermore, birds with brighter caps had larger clutches, suggesting that cap coloration may be an honest signal of parental quality in woodpeckers. Interestingly, more colourful individuals also occupied smaller territories, suggesting that territory size and territory quality may be inversely related in theMiddleSpottedWoodpecker.

Evidence that fathers, but not mothers, respond to mate and offspring coloration by favouring high-quality offspring

Parents have limited resources and, when simultaneously rearing multiple offspring, are expected to favour particularly valuable offspring. Because offspring characteristics like sex and ornamentation influence future reproductive value, these traits may influence parental favouritism. Parents in high-quality nest environments should prefer sons because sons show greater variability in reproductive potential than daughters. Moreover, when ornamental plumage coloration is condition dependent, parents should favour brighter over duller sons. We tested parental favouritism for fledgling-aged eastern bluebirds, Sialia sialis, a species in which fledglings display condition-dependent UV-blue coloration on their wings and tails. We simultaneously threatened two offspring with mock predators and recorded which offspring the parents defended. In accordance with the predicted value of offspring, fathers protected brighter over duller sons and protected sons over daughters when mated to high-quality mates. Mothers did not discriminate between sons and daughters or between brighter and duller sons. These results suggest that ornamental plumage coloration in juvenile bluebirds functions as a signal in father–offspring interactions. Paternal and maternal responses of eastern bluebirds to offspring characteristics may differ because of differences in parental allocation strategies or because males assess offspring quality based on variation in plumage coloration, a trait that also functions in competitive interactions among adult males.

A window on the past: male ornamental plumage reveals the quality of their early-life environment

It is well established that the expression of many ornamental traits is dependent on the current condition of the bearer. However, conditions experienced in early life are also known to be important for an individual's subsequent fitness and therefore, directly or indirectly, for the fitness of their mate. Specifically, a recent hypothesis suggests that sexually selected traits might be sensitive to conditions experienced during early-life development and thereby function as honest indicators of developmental history. Whether this applies to colourful male plumage, however, is largely unknown. We tested this idea with a field experiment by manipulating neonatal nutrition in a sexually dichromatic passerine, the hihi (Notymystis cincta). We found that carotenoid supplementation increased nestling plasma carotenoid concentration, which was in turn correlated with increased yellow saturation in male breeding plumage after moulting. We also found that the post-moult luminance (lightness) of the white ear-tufts tended to be reduced in males that had received an all-round nutritional supplement as nestlings. Black breeding plumage was not affected by neonatal nutritional treatment. Although the mechanisms that generate colourful plumage are evidently diverse, our results show that at least some parts of this display are accurate indicators of environmental conditions during development.

Haemosporidian parasites of a European passerine wintering in South Asia: diversity, mixed infections and effect on host condition

We studied haemosporidian parasites in the scarlet rosefinch Carpodacus erythrinus in a small isolated semicolony during an eight-year period using molecular methods of parasite detection. The scarlet rosefinch is an interesting model of parasite host species. It winters in South Asia which represents a rare exception among European passerines. Males express yellow to red carotenoid-based plumage ornament which is a good predictor of male reproductive success. In 240 blood samples originating from 199 adult individuals, the total parasite prevalence reached 60%. Prevalence varied among years from 36 to 81% in Haemoproteus, 8 to 22% in Plasmodium, and 0 to 14% in Leucocytozoon. Twenty parasite lineages were detected (Haemoproteus: 5 lineages, Plasmodium: 10 lineages, and Leucocytozoon: 5 lineages). Among them, the Haemoproteus ROFI2 lineage, which is a host-specific parasite lineage of the scarlet rosefinch, was the most frequently found. Parasite lineages showed varying degree of lineage specificity. While Haemoproteus lineages detected in the scarlet rosefinch have relatively narrow host breadth restricted mainly to Fringillidae family, Leucocytozoon and Plasmodium lineages generally showed wider host range. The presence of some parasite lineages hitherto detected in sedentary European passerines (SISKIN1, CCF3, BT2) or in Culicoides biting midges at the same locality (ROFI1) suggest local transmission. On the contrary, lineages LK05 and FANTAIL1 that were previously reported exclusively from Asian hosts imply parasite transmission at the scarlet rosefinch wintering sites in South Asia. Mixed infections were found in 17% of infected samples and comprised mainly the most frequent lineages. The pattern of concomitant infections seemed to be rather random and matched expected levels based on lineage frequencies. Between-year comparisons revealed that in a majority of the repeatedly captured individual hosts the infection status remained unchanged (individuals stayed uninfected or possessed the same parasite lineages). However, 16 gains and 8 losses of lineages were also reported. We have not found any effect of haemosporidians on male carotenoid ornament expression or host body mass.

Heterospecific exposure affects the development of secondary sexual traits in male zebra finches (Taeniopygia guttata)

In many animal species, social signals can affect the reproductive physiology and behaviour of conspecifics. In a few species that exhibit vocal learning, exposure to conspecific and sometimes heterospecific sounds can also influence their vocal development. Here we show that heterospecific exposure can affect the development of secondary sexual traits of male zebra finches (Taeniopygia guttata). In a first experiment, we trained young males with a passive playback of domesticated canary (Serinus canaria) song. Song playback enhanced the sexual development of the birds: they started to sing during the presentation of a video clip of a female earlier during development and exhibited secondary sexual plumage ornaments faster than males of the control group kept in silence. Two out of five birds exhibited clear evidence of imitation of canary song. In a second experiment, we raised young male finches with young male canaries in pairs until adulthood. Again, the live contact with a heterospecific companion affected the development of plumage ornaments in finches. We also observed some evidences of vocal convergence, both for calls and song. Moreover, young males of both species could recognize the call of their heterospecific companion when adults. These results suggest that heterospecific exposure can affect both the sexual and the vocal development of the zebra finch and can have long lasting effects.This article is part of a Special Issue entitled: insert SI title.

Multiple achromatic plumage ornaments signal to multiple receivers

We investigated whether achromatic plumage traits can act as multiple ornaments in an Arctic-breeding passerine, the snow bunting (Plectrophenax nivalis). Specifically, we examine whether multiple ornaments are providing multiple differing messages, are redundant, are unreliable signals of male quality, or are aimed at different receivers. We measured plumage reflectance and pigmentation patterns made conspicuous during male inter- and intrasexual displays that advertise different plumage regions. Our results indicate that although several aspects of male plumage may have redundant messages, different body regions appear aimed at different receivers. The wings of males—displayed primarily toward females during courtship—appear to indicate a pair’s future reproductive performance. Conversely, melanin-based plumage reflectance displayed during intrasexual threat displays provides information on territory features and a male’s capacity to defend it (i.e., territory size, territory quality, testosterone levels). Taken together, we suggest that snow buntings have multiple ornaments that provide information of differential importance in inter- versus intrasexual communication. This study demonstrates that achromatic plumage traits can serve in complex communication. Key wor ds: achromatic plumage, individual quality , multiple ornaments, multiple receivers, Plectrophenax nivalis. [Behav Ecol]

Long-lasting effects of yolk androgens on phenotype in the pied flycatcher (Ficedula hypoleuca)

The hormonal environment during early development, such as maternally derived androgens in bird eggs, shapes the development of the offspring in ways that may have important long-term consequences for phenotype and behavior and, ultimately, fitness. We studied the long-term effects of yolk androgens on several phenotypic and physiological traits in male and female pied flycatchers (Ficedula hypoleuca) by experimentally elevating yolk androgen levels and rearing birds in common-garden environment in captivity. We found that high yolk androgen levels increased the basal metabolic rates in both females and males in adulthood. High yolk androgen levels did not affect male melanin coloration or plumage ornaments, or timing or speed of moult in either sex. No effect of androgen treatment on cell-mediated or humoral immune response was found in either sex. Covariation among the measured phenotypic traits was further not altered by androgen treatment. Our results suggest that exposure to high androgen levels can have long-lasting effects on some offspring traits, but do not seem to lead to different phenotypes. Furthermore, the role of yolk androgens affecting sexually selected male traits in our study species seems to be minor. The fitness consequences of yolk androgen-induced higher metabolic rates remain to be studied.

Plumage reflectance signals dominance in Florida scrub-jay, Aphelocoma coerulescens, juveniles

Although sexual selection is undeniably important in the evolution and maintenance of ornamental traits, it remains an unsuitable explanation for juvenile ornamentation. Male–male competition has been important in the evolution of UV-blue plumage ornamentation in birds, suggesting that this type of plumage may signal status more generally. In the present study, we compared dominance status of juvenile Florida scrub-jays before and after experimental plumage manipulation and found that plumage reflectance signalled status. Male–male competition is important in maintaining UV-blue plumage ornamentation in adults of other species. However, our evidence shows that sexually dimorphic ornamentation can also mediate competition over food resources in juvenile Florida scrub-jays. Understanding status signalling among juveniles may yield insights into differences in fitness, especially in species with cohesive social systems where early interactions are likely to influence future competitions over resources or mates. ► We experimentally manipulated plumage reflectance of juvenile Florida scrub-jays. ► We compared dominance before and after plumage manipulation. ► We found that reducing plumage reflectance caused a reduction in dominance. ► We conclude that plumage reflectance signals status in juvenile Florida scrub-jays.

Multimodal sexual selection in a warbler: plumage and song are related to different fitness components

Many species possess multiple sexual signals that are produced in different modalities. Signals from different modalities may convey different or similar (redundant) information, or be intended for different receivers. We studied sexual selection on two such signals (plumage and song) in the common yellowthroat, Geothlypis trichas. We measured the coloration and size of both a carotenoid-based and a melanin-based plumage ornament and defined singing consistency as the lack of variability between repeated renditions of the same song. Overall, we found that plumage ornaments predicted within-pair mating success, while song consistency predicted extrapair success. Only plumage was related to male condition but, among experienced males, consistent singers had higher survival rates. Singing consistency was more variable than plumage, as performance varied across observations of the same male in relation to social context and date. Our results suggest that visual and acoustic signals in this species are most important in different sexual selection contexts.