Female Plumage Research Articles

Evolution of sexual dichromatism in relation to nesting habits in European passerines: a test of Wallace’s hypothesis

Wallace proposed in 1868 that natural rather than sexual selection could explain the striking differences in avian plumage dichromatism. Thus, he predicted that nesting habits, through their association with nest predation, could drive changes in sexual dichromatism by enabling females in cavity nesters to become as conspicuous as males, whereas Darwin (1871, The Descent of Man and Selection in Relation to Sex, John Murray, London) argued that sexual selection was the sole explanation for dichromatism. Sexual dichromatism is currently used as indicating the strength of sexual selection, and therefore testing Wallace's claim with modern phylogentically controlled methodologies is of prime interest for comparing the roles of natural and sexual selection in affecting the evolution of avian coloration. Here, we have related information on nest attendance, sexual dichromatism and nesting habits (open and cavity nesting) to male and female plumage conspicuousness in European passerines. Nest incubation attendance does not explain male or female plumage conspicuousness but nest type does. Moreover, although females of monochromatic and cavity nesting species are more conspicuous than females of other species, males of monochromatic and open nesting species are those with more cryptic plumage. Finally, analyses of character evolution suggest that changes in nesting habits influence the probability of changes in both dichromatism and plumage conspicuousness of males but do not significantly affect those in females. These results strongly suggest a role of nesting habits in the evolution of plumage conspicuousness of males, and a role for sexual selection also in females, both factors affecting the evolution of sexual dichromatism. We discuss our findings in relation to the debate that Darwin and Wallace maintained more than one century ago on the importance of natural and sexual selection in driving the evolution of plumage conspicuousness and sexual dichromatism in birds, and conclude that our results partly support the evolutionary scenarios envisaged by both extraordinary scientists.

Adaptive significance of permanent female mimicry in a bird of prey

Permanent female mimicry, in which adult males express a female phenotype, is known only from two bird species. A likely benefit of female mimicry is reduced intrasexual competition, allowing female-like males to access breeding resources while avoiding costly fights with typical territorial males. We tested this hypothesis in a population of marsh harriers Circus aeruginosus in which approximately 40 per cent of sexually mature males exhibit a permanent, i.e. lifelong, female plumage phenotype. Using simulated territorial intrusions, we measured aggressive responses of breeding males towards conspecific decoys of females, female-like males and typical males. We show that aggressive responses varied with both the type of decoys and the type of defending male. Typical males were aggressive towards typical male decoys more than they were towards female-like male decoys; female-like male decoys were attacked at a rate similar to that of female decoys. By contrast, female-like males tolerated male decoys (both typical and female-like) and directed their aggression towards female decoys. Thus, agonistic responses were intrasexual in typical males but intersexual in female-like males, indicating that the latter not only look like females but also behave like them when defending breeding resources. When intrasexual aggression is high, permanent female mimicry is arguably adaptive and could be seen as a permanent 'non-aggression pact' with other males.

Throat patch size and darkness covaries with testosterone in females of a sex-role reversed species

In male birds, size and blackness of melanin-based ornaments may signal competitive ability and quality. Testosterone is an important physiological mediator of these signals as these ornaments are testosterone dependent. Experimental manipulation of circulating testosterone in females suggests that similar mechanisms are at work in plumage ornaments of females. However, hormone manipulations typically result in testosterone levels that are far higher than physiological concentrations experienced by females. Thus, it remains unclear whether natural variation of testosterone levels would be sufficient to generate variation in female plumage coloration. A signaling value of plumage ornamentation is particularly important in females of sex-role reversed species, in which females are the more competitive sex. We examined the relationship between plumage coloration, body condition, and circulating testosterone in female barred buttonquails (Turnix suscitator), a classically polyandrous species. Even though female testosterone levels were low, they were strongly positively related with body condition and the size and blackness of the melanin throat patch. In males, no such relationships were apparent, suggesting that females may have an enhanced sensitivity for testosterone or its metabolites. These unique results reveal that a hormonal mechanism may be involved in sex-role reversal in this species.

Female plumage coloration is sensitive to the cost of reproduction. An experiment in blue tits

1. A growing number of studies suggest that female ornaments are linked to maternal quality and influence male mate choice. These findings challenge the traditional male-biased view of sexual selection and the hypothesis that female ornaments are the outcome of a genetic correlation with male ornaments. To further test the hypothesis that female traits have a function, it is now essential to investigate their honesty and to determine how signalling and reproduction interact in females. If female traits are honest indicators of quality, then they are likely to have a specific signalling function. 2. We investigated whether carry-over effects of reproduction might ensure the honesty of plumage colour signalling of a bird species with conspicuous UV-blue and yellow coloration, the blue tit Cyanistes caeruleus. Reproductive effort was manipulated by removing clutches, thereby forcing both sexes to reproduce twice and to raise chicks later in the breeding season when food is less abundant. In the year following this manipulation, we investigated the change in plumage in experimental and control males and females. The change was measured in the two putative feather ornaments, the UV-blue cap and the yellow breast, and another feather trait probably less likely to be sexually selected: the wing length. We also tested whether higher-quality females had their coloration less affected by the experiment. 3. We found that control but not manipulated males and females increased their signal towards UV. In addition, in the manipulated group, females that were able to lay more eggs had their UV-blue coloration less affected by the treatment. For yellow coloration, we found that manipulated yearlings but not manipulated adults decreased their yellow chroma in comparison with control. Lastly, our results show that the condition of the manipulated females tended to be positively correlated with yellow chroma. 4. These results show that the trade-offs between reproduction and signalling can ensure the honesty of conspicuous plumage traits in female and male blue tits. In addition, they suggest that female traits have the potential to evolve under sexual selection in this and other bird species.

Sources of distinctness of juvenile plumage in Western Palearctic passerines

Juveniles of many avian species possess a spotted or mottled body plumage that is visually distinct from the plumage of adults. In other species, however, juveniles fledge with a body plumage that is just a pale representation of adult female plumage. The reasons for this variation are poorly understood. Several hypotheses concerning social (parent‐offspring, adult‐juvenile, juvenile‐juvenile), ecological (predation risk) and physiological (costs of plumage development) implications of juvenile body plumage are presented in relation to predictions concerning associations with certain ecological and life-history attributes of avian species. In the present study, we conduct a phylogenetically corrected comparative analysis of Western Palearctic passerines looking for sources of variation in the incidence of distinct and adult-like juvenile body plumages. We scored plumages based on plates in the Handbook of the Birds of the Western Palearctic (Cramp & Perrins, 1988‐1994; Oxford University Press) (HBWP) and entered body mass, migratory habits, habitat, nestling diet, breeding dispersion, gregariousness, duration of the nestling period, type of nest, conspicuousness of female plumage, and sexual dimorphism as explanatory variables, as presented in HBWP, in phylogenetic generalized least square regression analyses. One-third of the species presented distinct juvenile body plumages, which lasted on average for the first 2 months of life. Body mass, conspicuousness of female plumage, migratory habits, and habitat were significantly associated with interspecific variation in distinctness of juvenile plumage, with smaller species, more conspicuous species, migrants, and species from forested habitats showing distinct juvenile plumages with higher frequency. The phylogenetic signal was moderately high. Assuming that conspicuous adult plumage is costlier to produce than distinct juvenile body plumage (pigments, conspicuousness), the need to acquire social status among juveniles before the winter may explain the more adult-like plumage in resident species because juveniles will probably compete with individuals that they may have known during their first months of life. On the other hand, migrant juveniles may compete with a different set of individuals in winter quarters and can use savings in resources necessary for developing adult-like plumages to improve migration capacity by allocating resources to other functions. The association with habitat could be related to juveniles in open habitats participating in more extended interactions with other juveniles than in forested habitats where lower visibility may reduce the capacity to detect or respond to signals from juvenile conspecifics. More studies on this possibly crucial life stage are needed. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 440‐454.

Analysis from avian visual perspective reveals plumage colour differences among females of capuchino seedeaters (Sporophila)

Females of the closely related capuchino seedeaters are difficult to distinguish from one another based on human visual perception of colouration and morphology. We examined plumage colour differences among females of four species, the tawny‐bellied seedeater Sporophila hypoxantha, the dark‐throated seedeater S. ruficollis, the rufous‐rumped seedeater S. hypochroma, and the chesnut‐bellied seedeater S. cinnamomea. Reflectance values were measured on museum skins, and interspecific differences were analyzed using the Vorobyev‐Osorio avian colour discrimination model. Interspecific distances in the colour space calculated by the model were considerably higher than the threshold for colour discrimination, indicating the presence of colour differences among species that should be detectable by birds. Colour differences between S. hypoxantha and the other three species were the largest. A Discriminant Function Analysis showed that UV‐wavelength was particularly important in species separation. Our results indicate that female plumage of these four species is considerably divergent in colour; these differences being imperceptible to the human eye, thus representing previously unknown morphological evolution in these species.

Nest Description of the Garden Emerald (Chlorostilbon assimilis) from Costa Rica

Abstract The Garden Emerald (Chlorostilbon assimilis) is endemic to southwestern Costa Rica and Panama, and knowledge about its' reproductive habits is limited. We describe the nest and nestlings of the Garden Emerald based on a nest found in La Amistad International Park. The nest was built on an anthropogenic substrate, and was similar to nests described for other emerald species. However, unlike other emeralds, the nest contained no lichens, mosses, or ferns. The nestlings resembled adult female plumage, similar to that for other nestling emerald species.

Increased plumage darkness of female Shiny Cowbirds Molothrus bonariensis in the subtropics: an adaptation to bacterial degradation?

The Shiny Cowbird Molothrus bonariensis is a sexually dichromatic species, in which males have blackish‐blue iridescence and females are dull brown. However, in some subtropical parts of its distribution, females show a plumage polymorphism that ranges from dull brown to dark brown and even black. Plumage melanization has been shown to protect feathers from bacterial degradation, decreasing the effects of harmful bacterial activity and thus plumage damage. In this study, we assessed whether bacterial feather‐degrading activity is acting as the selective force to increase darkness in the plumage of the female Shiny Cowbirds in Argentina. We compared the degradation of female Shiny Cowbird feathers belonging to different colour morphs when exposed to bacterial strains isolated from subtropical and temperate zones of its distribution, as well as to Bacillus licheniformis. We did not find differences in susceptibility to bacterial degradation between brown feathers and darker feathers. These results suggest that female plumage polymorphism in Shiny Cowbirds has not arisen as a defence against bacterial feather‐degrading activity.

Evidence that plumage bacteria influence feather coloration and body condition of eastern bluebirdsSialia sialis

Parasites influence the expression of secondary sexual traits and the health of infected individuals. We set out to test the influence of reputed exogenous parasites, plumage bacteria, including feather‐degrading bacteria (FDB), on secondary sexual characteristics and body condition of wild adult eastern bluebirdsSialia sialis. Previous work has shown that FDB alter the coloration of structurally‐colored bluebird feathersin vitro(Shawkey et al. 2007). In a correlational study of how bacteria affect birds in the wild, we found that female plumage got duller with increasing FDB intensity. Males tended to get brighter with increasing FDB intensity, but the relationship was not significant. We also found significant associations between plumage bacteria intensity and body condition, but, again, with gender‐based differences. Female body condition was negatively associated with plumage bacteria intensity, while male body condition was positively associated with plumage bacteria intensity. Interestingly, plumage bacteria intensity of males and females in nesting pairs was significantly positively correlated. We also report the highest prevalence of FDB measured in a wild bird population, with FDB detected on 67/68 (99%) of individuals. Further work is needed to fully understand the relationships between plumage bacteria and birds, but our data indicate that plumage bacteria may have sex‐dependent effects on multiple phenotypic traits.

Sexual dimorphic colours and female choice in the common kestrel

Common kestrels (Falco tinnunculus) exhibit pronounced sexual dimorphism in adult plumage colour. Since sexually dimorphic characteristics can be important in female choice and plumage colours represent signals of male quality in kestrels, we investigated whether female choice in captive common kestrels is influenced by the experimental manipulation of the grey colour of the male's tail and head feathers. In our experiment, females were allowed to choose between males with experimentally enhanced blue‐grey coloration (brighter than average) and control males (water treatment with no change in male colour patterns). We measured female preference as the frequency and duration of visits to each male (sitting on the perch or hanging on the partition wall). Overall, we did not detect any significant difference in frequency and duration of female visits between coloured and control males; however, individual females exhibited strong preferences for one of the two males, regardless of the colour of the male's tail and head. We briefly review hypotheses to explain why the plumage of the common kestrel is nevertheless sexually dimorphic.

A label of health: a previous immune challenge is reflected in the expression of a female plumage trait

Studies of ornaments as indicators of individual quality have hitherto focused on males, while studies of female ornaments have been almost absent. However, females within the same species may show large variation in both outer appearance and individual quality. We experimentally examined the effect of an immune challenge on the size of white fringes in the wing feathers produced one to two months after the challenge in female common eiders (Somateria mollissima), a long-lived sea duck. Immune-challenged females produced feathers with smaller white fringes, showing that a previous immune challenge may affect the expression of a plumage trait. We also report an unexpected difference in the expression of this white plumage trait within the immune-challenged group, related to the specific immune response against one of the injected antigens.

Plumage characteristics, reproductive investment and assortative mating in tree swallows Tachycineta bicolor

Elaborate ornamental plumage has been associated with various measures of individual quality in many species of birds. Male plumage characteristics, which have been relatively well studied, have been shown to reflect past reproductive investment, as well as the potential for reproductive investment in the current breeding attempt. In contrast, the signalling functions of female traits remain largely unexplored. In this study, we investigated the relationship between plumage attributes of breeding adult tree swallows and past reproductive investment, current reproductive investment and social mate pairing strategy. Both males and older females possess metallic green to metallic blue iridescent plumage on their dorsal surface, making this a suitable species for this type of investigation. We did not find any effects of past reproductive investment and success on the plumage attributes of returning breeders. In contrast, female plumage hue covaried with fledging success, and female plumage brightness was positively associated with mean clutch egg mass. In addition, we found that social pairs mated assortatively with respect to plumage brightness. We argue that since plumage characteristics vary with age in both male and female tree swallows, plumage attributes in this species are indicative of breeding experience and may be honest signals of quality. Positive assortative pairing could be the result of mutual mate choice or intra-sexual competition for nest sites by both males and females.

LACK OF ASSORTATIVE MATING FOR TAIL, BODY SIZE, OR CONDITION IN THE ELABORATE MONOMORPHIC TURQUOISE-BROWED MOTMOT (EUMOMOTA SUPERCILIOSA)

Elaborate male and female plumage can be maintained by mutual sexual selection and function as a mate-choice or status signal in both sexes. Both male and female Turquoise-browed Motmot (Eumomota superciliosa) have long tails that terminate in widened blue-and-black rackets that appear to hang, unattached, below the body of the bird. I tested whether mutual sexual selection maintains the Turquoise-browed Motmot's elaborate tail plumage by testing the prediction that mating occurs in an assortative manner for tail plumage. I also tested whether assortative mating occurs for body size, a potential measure of dominance, and for phenotypic condition, a measure of individual quality. Assortative mating was measured (1) within all pairs in the study population, (2) within newly formed pairs, and (3) within experimentally induced pairs that formed after removal of females from stable pairs. Assortative mating was not found for tail plumage, body size, or phenotypic condition in any of these samples. Therefore, there was no support for the “mutual sexual selection” hypothesis. I discuss the hypothesis that the tail is sexually selected in males only, and that natural selection accounts for the evolutionary maintenance of the elaborate female tail. Ausencia de Apareamiento Asociativo con Respecto a la Cola, el Tamaño Corporal o la Condición en Eumomota superciliosa

Female coloration indicates female reproductive capacity in blue tits

It is poorly understood whether female morphological and behavioural traits can be used as 'signals'. In particular, experimental tests of the hypothesis that female ornaments reflect quality are scarce. Here, we experimentally examine whether female plumage coloration might signal maternal quality in the blue tit, Cyanistes caeruleus by forcing half of the females breeding in our population to produce a replacement clutch. Using statistical models that controlled for the effects of male coloration, and the effects of age and condition of both parents, we found that carotenoid-based female coloration was positively linked to key proxies of bird lifetime reproductive success: clutch size, fledgling success and recruitment. Importantly, the relationships between maternal yellow carotenoid coloration and both clutch size and recruitment were stronger in the experimental group than in the control group, indicating that breeding females with higher values of yellow coloration were better able to handle the cost of producing a second clutch. Finally, UV-blue female coloration was positively linked to female survival and marginally linked to laying date. Taken together, these results show for the first time in a natural population that female coloration can indicate individual and maternal quality under natural and adverse reproductive conditions. They highlight the potential for the evolution of female ornamental traits through sexual selection.

Phenotypic plasticity in a conspicuous female plumage trait: information content and mating patterns

Explaining sexual ornamentation in the limiting sex, usually females, requires information on the proximate background of ornaments and their consequences for sexual selection. Phenotypic variation within individuals has received little attention in either of these research directions. We used 6 years of data to examine the information content and potential role of white wing patch size in female collared flycatchers, Ficedula albicollis. Female wing patch size differed among years. Yearling females had smaller wing patches than older females. The negative effect of original patch size on intraindividual patch size change was stronger in yearling than in older birds, which may reflect an age-dependent trade-off. Change in wing patch size was strongly positively related to the summer North Atlantic Oscillation (NAO) index. Clutch size laid in the previous breeding season had a negative effect on patch size change, but only in high-NAO years. Thus, the immediate effect of poor climate during moult apparently overrode the more indirect influence of reproductive effort. Two sexually selected ornaments of mates were unrelated to female wing patch size at the population level, but intraindividual changes of female wing patch size significantly predicted differences in wing patch size between mates obtained in the 2 years. Our data suggest that significant mating advantages to more ornamented females may not be detected from population-level mating patterns. Research on potential female ornaments should also pay more attention to age-dependent phenotypic plasticity, the trade-off between current and future ornament size and the costs of reproduction to apparent future attractiveness.

Iris colour as an indicator of age feature in female Brazilian tanagers (Passeriformes: Emberizidae) confirmed by a molecular sexing technique

The Brazilian tanager, Ramphocelus bresilius is an endemic species from Brazil that is sexually dimorphic in adult plumage. Young males are similar to adult and young females until their second year. Adults and young females are not distinguishable in plumage. We tested whether iris colour can be used to separate adult females from immature females. We used for the first time the molecular sexing technique based on CHD-genes to confirm the sex of the individuals classified as "female plumage with red iris", and to identify the sex of individuals classified as "female plumage and brown iris". The adult males were used as a positive control. DNA samples from 190 individuals were analysed. The sizes of the PCR products were identified as 350 base pairs (bp) for CHD-Z and 388 bp for CHD-W. We confirmed that adult females have a red iris and the young females a brown iris. We could also separate young males and females which present the same iris colour and plumage. Although there are indications that the iris colour can be used by birds to identify the adults in co-operative breeding species such as the Brazilian tanager, more behavioural data are required to understand the role of iris coloration in this species.

Melanin coloration in New World orioles I: carotenoid masking and pigment dichromatism in the orchard oriole complex

Carotenoids produce the brilliant red, orange, and yellow colors of many animals. However, melanin pigments can also confer some of these same hues. Because carotenoid and melanin colors are produced in different ways and may serve different signaling functions, either within or between species, it is important to establish whether one or both types of pigment are responsible for coloration. We have discovered what appears to be an evolutionary switch from carotenoid‐ to melanin‐based color in two sexually dichromatic New World orioles. Using a combination of reflectance spectrometry and chromatographic analyses of plumage pigments, we found that the chestnut plumage of adult male orchard oriolesIcterus spuriusis produced predominantly by phaeomelanins. Orchard oriole feathers also contain carotenoids, which appear to be masked by the high concentration of phaeomelanins. In contrast, both carotenoids and phaeomelanins appear to contribute to color in adult male Fuertes's oriolesI. fuertesi. Moreover, yellow yearling male and female plumage in both species is produced by carotenoids alone. The masking of carotenoids with phaeomelanins in orchard orioles is interesting in light of the signaling roles that carotenoids are thought to play. In addition, these plumage differences produce a unique case of age and sexual pigment dimorphism in orchard and Fuertes's orioles.

Unusual sex roles in a highly promiscuous parrot: the Greater Vasa ParrotCaracopsis vasa

We describe the unusual mating system of the Greater Vasa ParrotCaracopsis vasa. The dull black plumage of males and females is similar but females are significantly larger than males. Females are promiscuous and copulated with at least five different males. Copulations were either short (1–3 s) or very long (mean 35.9 min), and long copulations involved a copulatory tie facilitated by the male's enlarged cloacal protrusion. Multilocus DNA fingerprinting of 17 broods showed that all were of mixed paternity, and that some broods had three fathers. Males never visited nests directly, but during the incubation and chick‐rearing periods females came off the nest and were fed regurgitated fruit by multiple males. Four had band‐sharing coefficients that suggested they were unrelated. Males copulated with and provided food for several widely separated females simultaneously. During the chick‐rearing period females defended a territory around the nest from conspecific females, developed conspicuous orange skin on the head (through feather loss), and uttered loud, complex vocalizations that we refer to as ‘song’ from prominent perches near the nest. Males showed none of these traits. Females with high song rates attracted more males and as a result received more food than other females. Play‐back experiments in which female song rates were either increased or decreased, attracted more or fewer males respectively. We propose that female song, conspicuous head colour and territoriality have all evolved as a result of competition between females for the food provided by males. The selective pressures favouring this highly unusual breeding system in the Greater Vasa Parrot are unclear but some sort of ecological constraint, such as food availability, may be important.

Plumage Development and Molt in Long-Tailed Manakins (Chiroxiphia Linearis): Variation According to Sex and Age

AbstractLek-mating Long-tailed Manakins (Chiroxiphia linearis) exhibit an unusual pattern of delayed plumage maturation. Each year, males progress through a series of predefinitive plumages before attaining definitive plumage in their fifth calendar year. Females also exhibit variation in plumage coloration, with some females displaying male-like plumage characteristics. Using data from mist-net captures in northwest Costa Rica (n = 1,315) and museum specimens from throughout the range of Long-tailed Manakins (n = 585), we documented the plumage sequence progression of males, explored variation in female plumage, and described the timing of molt in this species. Males progressed through a series of age-specific predefinitive plumages, which enabled the accurate aging of predefinitive-plumaged males in the field; this predefinitive plumage sequence is the basis for age-related status-signaling in these males. Females tended to acquire red coloration in the crown as they aged. However, colorful plumage in females may be a byproduct of selection on bright male plumage. Females exhibited an early peak of molt activity from February to April, little molt from May through July, and a second, more pronounced peak of molt activity in October. By contrast, males in older predefinitive-plumage stages and males in definitive plumage exhibited comparable unimodal distributions in molt activity beginning in June and peaking between July and October. Our data are consistent with selective pressure to avoid the costs of molt-breeding overlap in females and older males. Our findings have important implications for social organization and signaling in Long- tailed Manakins, and for the evolution of delayed plumage maturation in birds.Desarrollo del Plumaje y Muda en Chiroxiphia linearis: Variación de Acuerdo al Sexo y la Edad

PLUMAGE DEVELOPMENT AND MOLT IN LONG-TAILED MANAKINS (CHIROXIPHIA LINEARIS): VARIATION ACCORDING TO SEX AND AGE

Lek-mating Long-tailed Manakins (Chiroxiphia linearis) exhibit an unusual pattern of delayed plumage maturation. Each year, males progress through a series of predefinitive plumages before attaining definitive plumage in their fifth calendar year. Females also exhibit variation in plumage coloration, with some females displaying male-like plumage characteristics. Using data from mist-net captures in northwest Costa Rica (n = 1,315) and museum specimens from throughout the range of Long-tailed Manakins (n = 585), we documented the plumage sequence progression of males, explored variation in female plumage, and described the timing of molt in this species. Males progressed through a series of age-specific predefinitive plumages, which enabled the accurate aging of predefinitive-plumaged males in the field; this predefinitive plumage sequence is the basis for age-related status-signaling in these males. Females tended to acquire red coloration in the crown as they aged. However, colorful plumage in females may be a byproduct of selection on bright male plumage. Females exhibited an early peak of molt activity from February to April, little molt from May through July, and a second, more pronounced peak of molt activity in October. By contrast, males in older predefinitive-plumage stages and males in definitive plumage exhibited comparable unimodal distributions in molt activity beginning in June and peaking between July and October. Our data are consistent with selective pressure to avoid the costs of molt-breeding overlap in females and older males. Our findings have important implications for social organization and signaling in Long- tailed Manakins, and for the evolution of delayed plumage maturation in birds. Desarrollo del Plumaje y Muda en Chiroxiphia linearis: Variación de Acuerdo al Sexo y la Edad