Male Plumage Research Articles

Testosterone Increase in Free-Living Young Blackcaps Sylvia atricapilla and Wood Warblers Phylloscopus sibilatrix During Post-Juvenile Molt with Possible Implications for Juvenile Dispersal

Testosterone may affect many aspects of passerine maturation including the expression of plumage coloration and spring migration in adult birds. However, how the testosterone level changes during bird development is not well known. We compared the testosterone profile during juvenile development of a sexually dimorphic species, the Blackcap Sylvia atricapilla, and a monomorphic species, the Wood Warbler Phylloscopus sibilatrix. We first tested whether testosterone influences plumage coloration and increases at the onset of molt in males of the dimorphic but not the monomorphic species. We found that a testosterone level increase occurred in both sexes and species during later stages of molt, and thus was not related to male plumage coloration. We also investigated whether the increase in testosterone level coincides with juvenile dispersal. If testosterone affects dispersal behavior in these species, both sexes should show an elevated testosterone level during dispersal, but this increase should occur earlier in the Wood Warbler, which disperses earlier than the Blackcap. In juvenile Blackcaps, the increase in testosterone level occurred on the 43-68th day after hatching, while in Wood Warblers it occurred on the 32-36th day (i.e. 11-32 days earlier). The increase in testosterone level coincided with the onset of the post-juvenile dispersal in both species. This study provides the first direct evidence of a testosterone level increase during juvenile development in two free-living migratory birds. While not correlated with species plumage coloration, this increase might affect other aspects of behavior, e.g. juvenile dispersal.

Female and male plumage color is linked to parental quality, pairing, and extrapair mating in a tropical passerine

AbstractSexual selection has been proposed to drive the evolution of elaborate phenotypic traits in males, which often confer success in competition or mating. However, in many species, both males and females display such traits. Studies examining how selection acts on both sexes are scarce. In this study, we investigated whether plumage ornamentation is sexually selected in females and males lovely fairy-wren Malurus amabilis, a cooperatively breeding songbird. We found that female and male plumage color was correlated with parental quality but not with individual quality or survival. We also found evidence of positive assortative mating based on plumage color. Microsatellite analyses of paternity indicated that the lovely fairy-wren has high levels of extrapair paternity (EPP), with 53% of offspring (in 58% of broods of 57% of females) resulting from extrapair (EP) mating. Female and male plumage color did not predict reproductive success or the proportion of EP offspring in their own nest, but less colorful males obtained higher EPP when paired with more colorful females and gained overall higher total paternity (own nest and other nests). We argue that plumage color may be under sex-specific selection, highlighting the importance of looking at both sexes in studies of sexual selection and ornament evolution. The current findings together with the previous study suggest that plumage color in female and male lovely fairy-wrens appears to be an honest signal relevant in both intrasexual and intersexual competition contexts.

Complex mosaic of sexual dichromatism and monochromatism in Pacific robins results from both gains and losses of elaborate coloration

Pacific robins exhibit one of the most complex range‐wide mosaics of sexual dichromatism and monochromatism. The evolutionary origins of this geographic mosaic remain poorly understood despite long‐standing interest from ornithologists, and its influential role in the development of Ernst Mayr's theories on speciation and the 'Biological Species Concept'. One factor limiting our understanding of the evolution of sexual plumage variation in Pacific robins is a lack of well‐resolved taxon boundaries and phylogenetic relationships in the group. Here, we use primarily historical museum specimens to obtain dense sampling of mtDNA, nuclear DNA, plumage color and morphometrics from all named taxa in the radiation in order to infer taxon boundaries and relationships. We use these data to test hypotheses about colonization history, plumage evolution and reduced island dichromatism. Our data show that the Pacific robin radiation comprises four distinct lineages, which warrant recognition as separate species – the previously recognized Norfolk robin P. multicolor and red‐capped robin P. goodenovii, and two new species we propose to name: ‘Solomon robin’ P. polymorpha Mayr, 1934 for the populations on Solomon and Bougainville Islands, and ‘Mayr's robin’ P. pusilla Peale, 1848 (in honor of Ernst Mayr's detailed work on the southwest Pacific robins) for the populations on Vanuatu, Fiji and Samoa. Our data suggest that the common ancestor of the entire Pacific robin radiation was most likely sexually dichromatic and that the radiation‐wide mosaic of sexual plumage color arose via repeated losses of elaborate plumage in males and gains of elaborate plumage in females on separate islands.

Hybridization between two recently diverged Neotropical passerines: The Pearly-bellied Seedeater Sporophila pileata, and the Copper Seedeater S. bouvreuil (Aves, Passeriformes, Thraupidae).

The small Neotropical finches called capuchinos are outstanding because they have experienced one of the most recent and explosive avian radiations ever documented for birds. Despite very low morphological and niche divergence among species, many of them are reproductively isolated when in sympatry due to strong sexual selection in plumage traits. However, a specific pair of mostly parapatric species, the Pearly-bellied, Sporophila pileata, and the Copper Seedeaters, S. bouvreuil, has confounded taxonomists because individuals with intermediate color patterns can be found. By analyzing diagnostic COI mtDNA sequences and adult male plumage we provide evidence for hybridization. Paternity tests using microsatellites also indicated that representatives with intermediate plumage pattern can be fertile. Our findings are consistent with the classification of S. bouvreuil and S. pileata as distinct taxa, but we demonstrate that the sexual selection mechanisms involved in the isolation of other reproductively sympatric capuchinos are not applicable to this pair of species, likely because of reduced barriers to mate recognition.

Natural and sexual selection shape the evolution of colour and conspicuousness in North American wood-warblers (Parulidae)

AbstractNatural and sexual selection drive colour evolution in animals. However, these different selective forces are often studied independently or without considering environmental variation. We evaluated the roles of natural and sexual selection together on colour evolution in 15 sympatric wood-warbler species, while considering the influence of variation in the light environment and visual background. We tested the influence of each selective pressure on male and female coloration and contrast against the background using avian visual models in phylogenetically controlled analyses. We found natural and sexual selection simultaneously driving cryptic and conspicuous plumage in males by acting on different body regions. For example, we found that ground-nesting species had males with conspicuous under-body plumage and cryptic upper-body plumage, showing how natural and sexual selection can drive colour evolution concordantly. We also found interesting relationships with female plumage, such as nest predation positively covarying with female contrast against the background, suggesting a cost to female conspicuousness. Our findings here showcase the complexity of selection on coloration and illustrate the importance of: (1) accounting for environmental variation when assessing how natural and sexual selection drive colour evolution; and (2) testing how multiple selection pressures are shaping colour diversity among species.

Pre- and post-copulatory traits working in concert: sexual dichromatism in passerines is associated with sperm morphology.

Passerine birds produce costly traits under intense sexual selection, including elaborate sexually dichromatic plumage and sperm morphologies, to compete for fertilizations. Plumage and sperm traits vary markedly among species, but it is unknown if this reflects a trade-off between pre- and post-copulatory investment under strong sexual selection producing negative trait covariance, or variation in the strength of sexual selection among species producing positive covariance. Using phylogenetic regression, we analysed datasets describing plumage and sperm morphological traits for 278 passerine species. We found a significant positive relationship between sperm midpiece length and male plumage elaboration and sexual dichromatism. We did not find a relationship between plumage elaboration and testes mass. Our results do not support a trade-off between plumage and sperm traits, but may be indicative of variance among species in the strength of sexual selection to produce both brightly coloured plumage and costly sperm traits.

Plumage iridescence is associated with distinct feather microbiota in a tropical passerine

Birds present a stunning diversity of plumage colors that have long fascinated evolutionary ecologists. Although plumage coloration is often linked to sexual selection, it may impact a number of physiological processes, including microbial resistance. At present, the degree to which differences between pigment-based vs. structural plumage coloration may affect the feather microbiota remains unanswered. Using quantitative PCR and DGGE profiling, we investigated feather microbial load, diversity and community structure among two allopatric subspecies of White-shouldered Fairywren, Malurus alboscapulatus that vary in expression of melanin-based vs. structural plumage coloration. We found that microbial load tended to be lower and feather microbial diversity was significantly higher in the plumage of black iridescent males, compared to black matte females and brown individuals. Moreover, black iridescent males had distinct feather microbial communities compared to black matte females and brown individuals. We suggest that distinctive nanostructure properties of iridescent male feathers or different investment in preening influence feather microbiota community composition and load. This study is the first to point to structural plumage coloration as a factor that may significantly regulate feather microbiota. Future work might explore fitness consequences and the role of microorganisms in the evolution of avian sexual dichromatism, with particular reference to iridescence.

Sex differences in song and plumage color do not evolve through sexual selection alone: new insights from recent research

Recent studies show that traits long recognized as sexually selected ornaments, such as the bright plumage colors and melodious vocal displays of songbirds, have evolved in surprising ways. Male plumage colors are generally more elaborate and diverse than female colors, and in many species only males sing, so both of these dimorphisms are widely presumed to have evolved through sexual selection on males. Yet, in multiple widely divergent songbird clades, sexual differences in colors and songs have evolved through losses in females rather than gains in males. Indeed, changes in female phenotypes may be the principle driving factor affecting such sexual dimorphisms in most songbirds. Sexual dichromatism and song dimorphism also tend to be associated with life history traits other than mating system, such as seasonal migration or territorial behaviors. Altogether, these recent findings support the view that sexual dimorphism is an evolutionary byproduct of multiple, different selection pressures acting on each sex rather than just sexual selection on males. Researchers should therefore be cautious to avoid assuming that dimorphism itself is a character solely reflecting levels of sexual selection in species.

Sexual selection predicts the rate and direction of colour divergence in a large avian radiation

Sexual selection is proposed to be a powerful driver of phenotypic evolution in animal systems. At macroevolutionary scales, sexual selection can theoretically drive both the rate and direction of phenotypic evolution, but this hypothesis remains contentious. Here, we find that differences in the rate and direction of plumage colour evolution are predicted by a proxy for sexual selection intensity (plumage dichromatism) in a large radiation of suboscine passerine birds (Tyrannida). We show that rates of plumage evolution are correlated between the sexes, but that sexual selection has a strong positive effect on male, but not female, interspecific divergence rates. Furthermore, we demonstrate that rapid male plumage divergence is biased towards carotenoid-based (red/yellow) colours widely assumed to represent honest sexual signals. Our results highlight the central role of sexual selection in driving avian colour divergence, and reveal the existence of convergent evolutionary responses of animal signalling traits under sexual selection.

Notes on nesting, territoriality and behaviour of broadbills (Eurylaimidae, Calyptomenidae) and pittas (Pittidae) in Tawau Hills Park, Sabah, Malaysian Borneo

Basic natural history and behavioural data are lacking for the majority of broadbills and pittas (Eurylaimidae, Calyptomenidae and Pittidae). We present a series of observations on these birds made during two visits to Tawau Hills Park, Sabah, Malaysia. During this period, we detected changes in temporal presence, detectability or vocal behaviour in various species. We also found habitat overlap on one ridgeline among four species of pitta. We observed a Green Broadbill Calyptomena viridis respond to our imitation of a call with a wing display. We discuss the social behaviour of Black-and-yellow Eurylaimus ochromalus and Banded Broadbills E. javanicus, including wing displays, territoriality, and in the first-named species, social groups comprising multiple adults. We also describe various nesting behaviours in these two species, including nest-construction techniques, incubation behaviour, nest defence, and the first described Banded Broadbill nest on Borneo. We found Dusky Broadbills Corydon sumatranus to frequently be the most conspicuous members of mixed-species flocks otherwise dominated by Cuculidae and Picidae. We analysed Black-crowned Pitta Erythropitta ussheri home range size and density in a colour-banded population, and found a potential hybrid or backcross with Garnet Pitta E. granatina. We observed that Blue-banded Pitta E. arquata make non-vocal sonations by striking their closed wings on their flanks, and discuss the immature male plumage of Blue-headed Pitta Hydrornis baudii.

Female and male plumage colour signals aggression in a dichromatic tropical songbird

Animal signals of competitive ability allow contests for limited resources to be settled without costly physical fights. Traits indicating competitive ability are diverse and span visual, acoustic or chemical modalities. Although animal signalling has been intensively studied, research has focused mainly on male traits. Little is known about the extent and functionality of competitive signals in females and whether there are sex differences in signal function. We studied whether plumage colour signals competitive ability in female and male lovely fairy-wrens, Malurus amabilis. In this species, both sexes sport elaborate but sexually dichromatic ornamental plumage. Using a mirror image stimulation test, we first assessed the relationship between male and female colour and agonistic behaviour, controlling for other physical, social and ecological variables. We then tested whether colourful plumage influenced aggressive response in both sexes by experimentally manipulating plumage colour and measuring individual responses to their mirror image. Females and males were more aggressive towards naturally less colourful reflections of the cheek patch in the mirror. However, when we manipulated plumage colour, both females and males responded more aggressively to experimentally increased cheek colour reflection in the mirror. Our findings suggest that plumage colour signals competitive ability in an aggressive context in both sexes and raises the possibility that signal reliability may be maintained by social interactions where individuals police and punish dishonest signals.

Minimum longevity and age-related male plumage in Darwin’s finches on Floreana Island

Several Darwin’s finch populations on the Galapagos Islands are declining and/or locally extinct. Acoustic surveys provide useful information about population size, but do not provide information on the age or morphology of birds. This study uses mist netting data collected during 2004–2016 on Floreana Island with the aim of evaluating minimum longevity in Darwin’s finches. The study species are the Small Tree Finch (Camarhynchus parvulus), hybrid tree finches (Camarhynchus spp.), Medium Tree Finch (Camarhynchus pauper), and Small Ground Finch (Geospiza fuliginosa). In total, 1032 Darwin’s finches were mist netted and 86 of 707 males and 14 of 325 females were recaptured across years. We used the proportion of black plumage to age males, and the age at first capture plus the number of years between recapture to estimate minimum longevity. Minimum longevity ranged from 12 to 15 years and was lowest in the critically endangered C. pauper (12 years). The average number of years between first capture and last recapture was significantly lower in females than in males. Because long-term mist netting provides information on age structure, recruitment and longevity in males and females, it should be a key component of effective conservation planning.

Experimental trait mismatches uncover specificity of evolutionary links between multiple signaling traits and their interactions in hummingbirds.

Many animal signals co-occur, and these signals may coevolve due to their interactive properties. Previous work has demonstrated ecological drivers of evolutionary relationships between signals and the environment, which leads to questions about why specific signal pairs evolved among species that possess multiple signals. We asked whether the coloration of different species was optimized for presentation with its natural behavioral display. We investigated this in "bee" hummingbirds, where males exhibit angle-dependent structurally-colored plumage and a stereotyped courtship (shuttle) display, by experimentally creating mismatches between the behavior and plumage of five species and quantifying how these mismatches influenced male color appearance during a display. Specifically, we photographed the plumage from a given species as we moved its feathers through the position-and-orientation-specific courtship display path of other species and quantified the resulting color appearance during the display in order to compare the mismatched color appearance to each species' natural color appearance. We found that mismatches significantly altered display flashiness (% change in coloration during displays) compared to the natural plumage-behavior pairings, and that such departures in flashiness were predicted by differences in shuttle behaviors alone. These results illustrate a tight evolutionary relationship between shuttle displays and color flashiness in these hummingbirds. Further, we found that interspecific variation in male plumage, behavior, and natural color appearance predicted deviations between natural and mismatched flashy color appearance. Altogether, our work provides a new method for testing signal coevolution and highlights the complex evolutionary relationships between multiple signals and their interactions.

Taxonomy of the White-browed Shortwing (Brachypteryx montana) complex on mainland Asia and Taiwan: an integrative approach supports recognition of three instead of one species

BackgroundThe White-browed Shortwing (Brachypteryx montana) is widespread from the central Himalayas to the southeast Chinese mainland and the island of Taiwan, the Philippines and Indonesia. Multiple subspecies are recognised, and several of these have recently been suggested to be treated as separate species based on differences in morphology and songs.MethodsWe here analyse plumage, morphometrics, songs, two mitochondrial and two nuclear markers, and geographical distributions of the two mainland Asian taxa B. m. cruralis and B. m. sinensis and the Taiwanese B. m. goodfellowi.ResultsWe conclude that these differ congruently in morphology, songs and DNA. Male B. m. goodfellowi is the most divergent in plumage (sexually monomorphic, unlike the two others; male similar to female), and B. m. cruralis and B. m. sinensis differ in male plumage maturation. The song of B. m. cruralis is strongly divergent from the others, whereas the songs of B. m. sinensis and B. m. goodfellowi are more similar to each other. Brachypteryx m. sinensis and B. m. goodfellowi are sisters, with an estimated divergence time 4.1 million years ago (mya; 95% highest posterior distribution [HPD] 2.8–5.5 mya), and B. m. cruralis separated from these two 5.8 mya (95% HPD 4.1–7.5 mya). We also report notable range extensions of B. m. sinensis as well as sympatry between this taxon and B. m. cruralis in Sichuan Province, China. Brachypteryx m. montana from Java is found to be more closely related to Lesser Shortwing (B. leucophris) and Rusty-bellied Shortwing (B. hyperythra) than to the mainland Asian and Taiwanese taxa.ConclusionOur data support a recent proposal to treat the three mainland Asian and Taiwanese taxa as three species, separate from B. montana sensu stricto: B. cruralis (central Himalayas to south central China and south Vietnam), B. sinensis (north central to southeastern part of mainland China) and B. goodfellowi (Taiwan Island).

Baseline corticosterone does not reflect iridescent plumage traits in female tree swallows

The production of high quality secondary sexual traits can be constrained by trade-offs in the allocation of energy and nutrients with other metabolic activities, and is mediated by physiological processes. In birds, the factors influencing male plumage quality have been well studied; however, factors affecting female plumage quality are poorly understood. Furthermore, it remains uncertain which physiological traits mediate the relationship between body condition and ornaments. In this three-year study of after-second-year female tree swallows (Tachycineta bicolor), we investigated (1) the relationship between baseline corticosterone near the end of the brood-rearing period (CORTBR) and feather colour characteristics (hue, saturation, brightness) the following year, and (2) the relationship between baseline corticosterone measured during incubation (CORTI) and brood rearing (CORTBR), and feather colour in the same year. To control for reproductive effort, we included reproductive parameters as covariates in all analyses. In this first study between CORT and the plumage colour characteristics of a species bearing iridescent feathers, we did not find any relationship between CORTBR and the colour of subsequently-produced feathers, nor did we find any relationship between CORT and the colour of feathers displayed during that breeding season. If CORT levels at the end of breeding carry over to influence the immediately subsequent moult period as we expect, our results generally indicate that structural plumage quality may not be as sensitive to circulating CORT levels compared to carotenoid-based colouration. Future studies, particularly those employing experimental manipulations of CORT during moult in species with iridescent traits, are necessary to fully determine the role glucocorticoids play in mediating the quality of secondary sexual characteristics.

A striking, critically endangered, new species of hillstar (Trochilidae: Oreotrochilus) from the southwestern Andes of Ecuador

ABSTRACT We describe a new species of the genus Oreotrochilus from the southwestern Andes of Ecuador. The new species is most similar in adult male plumage to O. stolzmanni and O. chimborazo. However, male and female show unique combinations of plumage characters that are likely to act as social signals. Phylogenetic analyses based on mitochondrial DNA indicate that this new taxon is closely related to O. stolzmanni and O. melanogaster, whereas genetic distances and preliminary comparisons of vocalizations suggest a sister relationship with O. stolzmanni. The geographic distribution of the new species seems to be restricted to cordillera Chilla-Tioloma-Fierro Urcu, in the southwestern highlands of Ecuador, an area historically poorly explored by ornithologists. Thus, based on its restricted distribution, apparently low population size, and lack of protection of its habitat, we evaluate it as critically endangered.

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.

Mutations in the melanocortin-1 receptor (MC1R) gene have no influence on the distinct patterns of melanic plumage found in the manakins of the genus Antilophia (Aves: Pipridae).

The melanocortin-1 receptor gene is the most widely-used marker for the investigation of the genetic determination of melanic plumage patterns. Studies of a number of wild bird species have shown an association between non-synonymous mutations of the MC1R gene and the presence of melanic variants. The genus Antilophia (Pipridae) includes only two manakin species (A. galeata and A. bokermanni), which are distinguished primarily by the differences in the pattern of melanic coloration of the plumage of the mantle in the adult males. In A. galeata, this plumage is black, while in A. bokermanni, it is predominantly white. This study investigates the possible association between mutations of the MC1R marker and the variation in plumage coloration observed in the two species. The MC1R sequences of the two species was analyzed, and the observed nucleotide variation was compared. Six polymorphic sites were identified, representing seven distinct genotypes. Five of these polymorphic mutations were non-synonymous, but were not related to the different phenotypes. Neutral evolution and the absence of any systematic association between the variants of the MC1R and plumage coloration in the Antilophia species indicate that alternative mechanisms regulate the expression of the coloration of the plumage in the adult males.

Rufous Common Cuckoo chicks are not always female

The Common Cuckoo (hereafter Cuckoo) shows two adult plumage morphs—adult male plumage is grey and adult females are either grey or, less frequently, rufous. The situation is less clear in juveniles, as both sexes exhibit variable proportions of grey and rufous colour. We thus describe the patterns related to sex-specific plumage colour variation in a central European Cuckoo population. We genetically determined the sex of 91 Cuckoo chicks and using visual classification of photographs we scored juvenile plumage colouration of individual chicks into five classes based upon the increasing proportion of rufous colour on feathers. To verify these scores, we sampled chick feathers and quantified the proportion of rufous colour of individual feathers by digital image analysis. We found that juvenile females had a higher proportion of rufous colour of feathers than juvenile males. However, the difference was marginally non-significant based on visual inspection alone, and some male chicks even showed intensively rufous plumage like those of juvenile females. In contrast, we captured only grey adult males (n = 37), while five out of 20 adult females were rufous. The rufous colour of Cuckoo feathers considerably differed from the grey colour and the difference seemed to be larger in adults than in juveniles. We show that chicks, unlike adult females, cannot be visually assigned to either of the adult morphs. Therefore, we encourage further investigation of Cuckoo plumage colouration across the species’ range to examine the process of plumage maturation. A detailed genetic analysis is necessary to understand the origin of Cuckoo feather colouration.

Sexual Selection Mechanisms for Male Plumage Ornaments in Japanese Barn Swallows

The Barn Swallow Hirundo rustica is a model species for studying sexual selection. After its long tail was shown to be the target of female mate choice, its plumage characteristics, including the red throat patch and the size of white tail spots in addition to tail length, have attracted the attention of many researchers. Although there have been several reviews of sexual selection in this species, these reviews have focused mostly on general patterns of sexual selection within and among subspecies, rather than on the functional differences between plumage characteristics in sexual selection. Here, I review mechanisms of sexual selection for male plumage ornaments in Japanese Barn Swallows H. r. gutturalis. In the last decade, studies of sexual selection in Japanese Barn Swallows have focussed on tail length, the size of the white tail spots, and the size and coloration of red throat patches. Each ornamental trait appears to be involved in different sexual selection mechanisms: the size of the white tail spots is important for several mechanisms of intersexual selection, including the Darwin-Fisher mechanism and differential allocation, whereas throat coloration may be intra-sexually selected, at least in part. The size of the throat patch, in contrast, is the target of an intersexual selection mechanism, differential access, indicating a partially independent evolution of the two components of the red throat patch. Multiple mechanisms of sexual selection would explain the multiple ornaments and their geographical variation, and even the probability of speciation and extinction.