Covariation between bill colouration and fitness components in a nocturnal bird

Vividly coloured chromatic signals play a key role in social and sexual signalling in diurnal birds, but their role is considered negligible in favour of achromatic (i.e. white, pure grey and black colourations) signals in nocturnal species. Here we studied colour variation and potential signalling of the yellow bill – a trait functioning as sexual signal in diurnal raptors during the breeding season in females of a truly nocturnal raptor, the little owl Athene noctua. We found that yellow‐red chroma of the bill was highly variable between individuals and positively correlated with female fitness prospects (i.e. brood size at fledging). In addition, we found that females with brighter bills were larger in size and produced owlets with a higher mass at fledging. This study suggests that yellow bill colouration in female little owls may potentially play a role in sexual signalling and may constitute the first evidence of chromatic colour signalling in a nocturnal bird.     

Plumage colour in nestling blue tits: sexual dichromatism,condition dependence and genetic effects

Sexual-selection theory assumes that there are costs associated with ornamental plumage coloration. While pigment-based ornaments have repeatedly been shown to be condition dependent, this has been more difficult to demonstrate for structural colours. We present evidence for condition dependence of both types of plumage colour in nestling blue tits (Parus caeruleus). Using reflectance spectrometry, we show that blue tit nestlings are sexually dichromatic, with males having more chromatic (more 'saturated') and ultraviolet (UV)-shifted tail coloration and more chromatic yellow breast coloration. The sexual dimorphism in nestling tail coloration is qualitatively similar to that of chick-feeding adults from the same population. By contrast, the breast plumage of adult birds is not sexually dichromatic in terms of chroma. In nestlings, the chroma of both tail and breast feathers is positively associated with condition (body mass on day 14). The UV/blue hue of the tail feathers is influenced by paternally inherited genes, as indicated by a maternal half-sibling comparison. We conclude that the expression of both carotenoid-based and structural coloration seems to be condition dependent in blue tit nestlings, and that there are additional genetic effects on the hue of the UV/blue tail feathers. The signalling or other functions of sexual dichromatism in nestlings remain obscure. Our study shows that nestling blue tits are suitable model organisms for the study of ontogenetic costs and heritability of both carotenoid-based and structural colour in birds.     

The role of colour in signalling and male choice in the agamid lizard Ctenophorus ornatus

Bright coloration and complex visual displays are frequent and well described in many lizard families. Reflectance spectrometry which extends into the ultraviolet (UV) allows measurement of such coloration independent of our visual system. We examined the role of colour in signalling and mate choice in the agamid lizard Ctenophorus ornatus. We found that throat reflectance strongly contrasted against the granite background of the lizards' habitat. The throat may act as a signal via the head-bobbing and push-up displays of C. ornatus. Dorsal coloration provided camouflage against the granite background, particularly in females. C. ornatus was sexually dichromatic for all traits examined including throat UV reflectance which is beyond human visual perception. Female throats were highly variable in spectral reflectance and males preferred females with higher throat chroma between 370 and 400 nm. However, female throat UV chroma is strongly correlated to both throat brightness and chest UV chroma and males may choose females on a combination of these colour variables. There was no evidence that female throat or chest coloration was an indicator of female quality. However, female brightness significantly predicted a female's laying date and, thus, may signal receptivity. One function of visual display in this species appears to be intersexual signalling, resulting in male choice of females.     

Informative content of melanin‐based plumage colour in adult Eurasian kestrels

Covariation between melanin‐based colorations and other phenotypic attributes has been rarely measured simultaneously in males and females. Such covariations and mechanisms mediating them have crucial importance determining the signalling function of these coloured ornaments in the two sexes. We examined the role of four melanin‐based coloured plumage patches as indicators of quality in both males and females of the sexually dimorphic and dichromatic Eurasian kestrel Falco tinnunculus. Previous kestrel studies have focused on the size of melanin plumage patches in either males or females as indicators of individual quality. Here, we used spectrophotometric measurements of three plumage patches and the size of another plumage patch to investigate the information content of multiple plumage colour traits in male and female kestrels. We found that females with bright plumage in the head and with high UV chroma in black parts of the rump showed good body condition and innate immunity. In addition, laying date was significantly explained by the intensity of the brown in the head of females. Meanwhile, in males we only found that individuals with greyer rumps showed better innate immunity. Altogether, our results indicate that, irrespective of the mechanism promoting covariation between coloration and individual quality, melanin‐based coloration can inform on individual quality in adult kestrels.     

Interspecific variation in plumage colour among birds: species recognition or light environment?

Abstract The traditional explanation for interspecific plumage colour variation in birds is that colour differences between species are adaptations to minimize the risk of hybridization. Under this explanation, colour differences between closely related species of birds represent reproductive character displacement. An alternative explanation is that interspecific variation in plumage colour is an adaptive response to variation in light environments across habitats. Under this explanation, differences in colour between closely related species are a product of selection on signal efficiency. We use a comparative approach to examine these two hypotheses, testing the effects of sympatry and habitat use, respectively, on divergence in male plumage colour. Contrary to the prediction of the Species Isolation Hypothesis, we find no evidence that sympatric pairs of species are consistently more divergent in coloration than are allopatric pairs of species. However, in agreement with the Light Environment Hypothesis, we find significant associations between plumage coloration and habitat use. All of these results remain qualitatively unchanged irrespective of the statistical methodology used to compare reflectance spectra, the body regions used in the analyses, or the exclusion of areas of plumage not used in sexual displays. Our results suggest that, in general, interspecific variation in plumage colour among birds is more strongly influenced by the signalling environment than by the risk of hybridization.     

Revealing the colourful side of birds: spatial distribution of conspicuous plumage colours on the body of Australian birds

In many species of birds, different body parts often display very different colours. This spatial distribution of coloured plumage patches may be determined, among other factors, by the balance between being cryptic to predators, and conspicuous to intended receivers. If this is the case, ventral and anterior body parts in birds – which are less visible to predators but more prominent to conspecifics – should present more conspicuous and sexually dichromatic plumage colours. Here, I test these predictions using reflectance spectrometric measurements of standardised plumage patches across males and females for nearly an entire avifauna (Australian landbirds, n = 538 species). My data show that, as predicted, conspicuous and sexually dichromatic colours are mainly located near the head, while the plumage of the back is the most cryptic. One clear exception to this pattern is the conspicuous rump coloration. In many species, this patch can be concealed by wings, and therefore exposed only when necessary. In addition, conspicuous rump coloration could deflect or confuse predators in case of attack. However, there is considerable variation across species, and this makes position on the body a very poor predictor of plumage elaboration (R² < 0.02). Future studies should try to determine whether differences between species in the distribution of colours across the plumage are due to variation in ecological factors (predation risk, habitat, etc.).     

Comparing plumage colour measurements obtained directly from live birds and from collected feathers: the case of the great tit Parus major

Birds frequently display a colourful plumage which is important both in inter and intraespecific communication, and either in sexual and social contexts. In last years some methodologies have been developed to, analyse plumage coloration, but the use of the spectrometers has been particularly important for UV range. Measurement of plumage coloration with the spectrometer may be taken directly on the bird or, alternatively by collecting some feathers and measuring them later in the laboratory. However, few is known about the reliability of measures obtained from feathers and whether these are really representative of plumage coloration. We tested this assumption analysing measurements of carotenoids-based coloration components (lightness, chroma and hue) and lutein peak of the yellow breast of the great tit Parus major. We used two spectrometers (Ocean optics and Minolta) which calculate differently the colour components. Our results showed that direct measurement of bird was highly repeatable to determine lightness, chroma and hue for both spectrometers. Similar results we found for collected feathers procedure for both devices. Collected feathers provided high representative measurements of colour values with Minolta spectrometer. Lightness was highly repeatable when we used Ocean optic spectrometer, but chroma and hue were moderate. Lutein peak was also highly repeatable in all cases. The number of feathers used to measure plumage coloration in collected feathers procedure strongly influenced values of colour plumage variables. In general, values of lightness, chroma and hue stabilised when more than 10–15 feathers were used although we found slight differences between spectrometers. However, only four feathers were needed for lutein peak. Thus, our results stress the need to use a minimum number of feathers in measuring plumage coloration from collected feathers.     

Seasonal Changes in Colour: A Comparison of Structural,Melanin- and Carotenoid-Based Plumage Colours

Background

Plumage coloration is important for bird communication, most notably in sexual signalling. Colour is often considered a good quality indicator, and the expression of exaggerated colours may depend on individual condition during moult. After moult, plumage coloration has been deemed fixed due to the fact that feathers are dead structures. Still, many plumage colours change after moult, although whether this affects signalling has not been sufficiently assessed.

Methodology/Principal Findings

We studied changes in coloration after moult in four passerine birds (robin, Erithacus rubecula; blackbird, Turdus merula; blue tit, Cyanistes caeruleus; and great tit, Parus major) displaying various coloration types (melanin-, carotenoid-based and structural). Birds were caught regularly during three years to measure plumage reflectance. We used models of avian colour vision to derive two variables, one describing chromatic and the other achromatic variation over the year that can be compared in magnitude among different colour types. All studied plumage patches but one (yellow breast of the blue tit) showed significant chromatic changes over the year, although these were smaller than for a typical dynamic trait (bill colour). Overall, structural colours showed a reduction in relative reflectance at shorter wavelengths, carotenoid-based colours the opposite pattern, while no general pattern was found for melanin-based colours. Achromatic changes were also common, but there were no consistent patterns of change for the different types of colours.

Conclusions/Significance

Changes of plumage coloration independent of moult are probably widespread; they should be perceivable by birds and have the potential to affect colour signalling.     

Seasonality of carotenoid‐based plumage coloration: modelling wavelength‐specific change through spectral reconstruction

Plumage coloration has provided important model systems for research on signal expression. Whilst it had previously been assumed that moulting provided the only mechanism to change plumage coloration, recent studies have shown plumage colours to be seasonally dynamic, with implications both for the quantification of expression and for any signalling role. However, the mechanistic processes underlying such change remain uncertain. Here, we describe within‐moult shifts in expression of a carotenoid‐based colour trait – the yellow ventral plumage of the great tit Parus major – over a nine‐month timespan. We report that plumage chromaticity (‘colour’) – but not achromaticity (‘brightness’) – exhibits a marked seasonal decline, independent of sex, age or body condition, and at a constant rate across twelve environmentally heterogeneous plots within our study site. To gain a greater understanding of the mechanisms underlying this change we employed a spectral reconstruction approach, that generates predicted spectra for any timepoint within the sampling period. By comparing spectra for both early and late in the moult we show that the seasonal decline in chromaticity is driven by both a marked reduction in ultraviolet (UV) reflectance and, to a lesser extent, loss of active carotenoid pigments. Thus, our study shows that seasonal loss of chromaticity in the great tit is driven by altered reflectance primarily in the UV section of the spectrum, a finding made possible by the use of spectral compartmentalisation and multi‐parallel modelling to produce reconstructed spectra. Whether change in plumage coloration influences signal function will depend on the dynamics of the signalling system but it could clearly inflate patterns such as assortative mating and should be considered in studies of colour expression.     

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.     

Sex and age differences in reflectance and biochemistry of carotenoid‐based colour variation in the great tit Parus major

The plumage coloration in great tits (Parus major) is the subject of much behavioural and ecophysiological research, yet there is a lack of analyses of the natural colour variation and its mechanisms. We used reflectance spectrometry and high‐performance liquid chromatography to explore individual, sexual and age‐related variation in carotenoid coloration and pigmentation, paramount to the often presumed, but rarely substantiated, costs and ‘honesty’ of carotenoid displays. In adults, we found that sex was the strongest predictor of ‘brightness’ (higher in males) and of ‘hue’ (longer wavelength in females). There was no sex difference in ‘carotenoid chroma’ or carotenoid content of feathers which also was unrelated to adult age (1 or 2+ years) and condition. Similar patterns were revealed for nestlings. Regarding the biochemical ‘signal content’, ‘carotenoid chroma’, but not ‘hue’, was significantly related to the carotenoid content (lutein and zeaxanthin) of feathers. These results refute the previously assumed exaggeration of carotenoid pigmentation in male great tits, and question the condition‐dependence of carotenoid coloration in this species. However, the sexual dimorphism in total reflectance or ‘brightness’, most likely due to melanins rather than carotenoids, may have implications for signalling or other adaptive explanations that need to be explored. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 758–765.     

Structural plumage colour and parasites in satin bowerbirds Ptilonorhynchus violaceus: implications for sexual selection

We investigated whether variation in structural plumage coloration in satin bowerbirds, Ptilonorhynchus violaceus , could reveal the intensity of infection from parasites, as predicted from models of parasite-mediated sexual selection (PMSS). To do this, we captured adult male, female, and juvenile male satin bowerbirds in Queensland, Australia, and objectively measured individual plumage reflectance from four body regions using a spectrometer. We quantified both ectoparasite load and the intensity of infection from blood parasites. In iridescent blue adult males, plumage reflectance is unimodal, with a single peak in the ultraviolet, while in greenish females and juveniles, plumage reflectance is bimodal, with peaks in both the ultraviolet and green portions of the spectrum. In adult males, the intensity of infection from blood parasites was best predicted by plumage brightness (total reflectance), with brighter males having fewer parasites. Similarly, juvenile males exhibiting greater UV chroma (proportion of reflectance in the UV) had fewer blood parasites. Our findings support a key prediction of PMSS models and provide the first evidence that a structural colour ornament can signal the intensity of infection from blood parasites.     

Selection and inheritance of sexually dimorphic juvenile plumage coloration

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

Plumage coloration can be perceived as a multiple condition‐dependent signal by Great Tits Parus major

The evolution of multiple signals can be explained because they enhance the perception of a general message by recipients. Plumage coloration frequently acts as a condition‐dependent signal, so that species displaying different colour patches have the potential to transmit information on condition through a multiple signalling system. The Great Tit Parus major exhibits plumage colour patches generated by the main types of colour production, some of which, particularly those based on melanins and carotenoids, are known to be related to body condition. However, the colour expressions of all of the different colour types have never been investigated simultaneously to determine whether they function as multiple signals of condition. In addition, visual perception models have never been applied to a multiple signalling system in a wild population of birds. Here I present information that links body condition with the colour expression of almost all of the different plumage patches of male Great Tits captured during the winter. Birds in better condition had greater reflectance values at short wavelengths in all plumage patches, and this was especially so in the white (i.e. structural) colour of the cheeks. Plumage colour characteristics were calculated by means of avian visual models, suggesting that Great Tits have the capacity to perceive information contained in the plumage coloration of conspecifics. These results show that short‐wavelength reflectance has great potential to transmit biologically significant information on the body condition of birds, even in achromatic plumage patches.     

Brightness variability in the white badge of the eagle owl Bubo bubo

The application of modern spectrometry to the study of avian colour variability has revealed ignored patterns of colour variation such as male‐biased sexual dichromatism and seasonal variability in the plumage. However, the variation in the achromatic property of such traits, that is in the total light reflectance of the spectrum (i.e., brightness), has commonly been overlooked. The evolution of signals based on brightness should be favoured in those species that are active when light is scarce, i.e. at dawn and dusk. The eagle owl Bubo bubo is monogamous and apparently monomorphic in plumage‐coloration. In this species, sexual and territorial call behaviour is mainly performed at dawn and dusk, during which a white patch on the throat is repeatedly exposed at each call. We measured the total light reflectance of the feathers of this badge in 39 eagle owl specimens from museum collections. We found seasonal variability and sexual dichromatism in the brightness of the plumage badge. The total reflectance of this trait peaked during the territorial‐mating period. Moreover, females showed higher values of brightness than males, in agreement with the reversed body size dimorphism present in this and many other raptor species. Finally, female but not male body size was positively correlated with white badge reflectance.     

Reproductive correlates of plumage coloration of female Mountain Bluebirds

Many studies have shown that the plumage coloration of male birds can act as an honest signal of quality, indicating benefits that a female could gain from pairing with a specific male. In some species, females also display ornamental plumage, but less is known about the function and potential adaptive significance of female coloration because most research has focused on male coloration. Male Mountain Bluebirds (Sialia currucoides) display full body, ultraviolet (UV)‐blue plumage, whereas female plumage is more subdued, with blue color focused on the rump, wing, and tail. During the 2011 and 2012 breeding seasons (May–July) near Kamloops, BC, Canada, we examined coloration of the rump and tail of female Mountain Bluebirds to determine if their plumage could act as an indicator of direct reproductive benefits (e.g., enhanced parental care or reproductive success) to potential mates. We found no relationship between female plumage coloration and either provisioning rate or fledging success. However, female coloration varied with age, with after‐second‐year (ASY) females having brighter, more UV‐blue tail feathers than second‐year (SY) females. In addition, ASY females with brighter, more UV‐blue tails had larger clutches. We also observed positive assortative mating by tarsus length. Because previous work with other species suggests that female body size may be a good predictor of breeding success, males could potentially benefit from pairing with larger females. However, reproductive success did not vary with female size in our study. Although our evidence that structural plumage coloration of female Mountain Bluebirds is a signal of direct reproductive benefits for males (e.g., higher reproductive success) is limited, our results (i.e., ASY females with brighter tails than SY females, and ASY females with brighter tails having larger clutches) do suggest the potential for sexual selection to act on female coloration.     

Nanostructure predicts intraspecific variation in ultraviolet-blue plumage colour

Evidence suggests that structural plumage colour can be an honest signal of individual quality, but the mechanisms responsible for the variation in expression of structural coloration within a species have not been identified. We used full-spectrum spectrometry and transmission electron microscopy to investigate the effect of variation in the nanostructure of the spongy layer on expression of structural ultraviolet (UV)-blue coloration in eastern bluebird (Sialia sialis) feathers. Fourier analysis revealed that feather nanostructure was highly organized but did not accurately predict variation in hue. Within the spongy layer of feather barbs, the number of circular keratin rods significantly predicted UV-violet chroma, whereas the standard error of the diameter of these rods significantly predicted spectral saturation. These observations show that the precision of nanostructural arrangement determines some colour variation in feathers.     

Carotenoid-based coloration correlates with the hatching date of Blue Tit Cyanistes caeruleus nestlings

Carotenoid-based coloration occurs predominantly in adult birds, yet in some species from the family Paridae, this trait is also present at the nestling stage. One of the factors proposed to affect the expression of this trait in immature birds is hatching date. Here, using the avian tetrahedral colour space model, we examined the influence of hatching date on the breast carotenoid-based plumage coloration of the Blue Tit Cyanistes caeruleus nestlings. Because Blue Tits are sexually dichromatic, we also investigated the potential interaction between hatching date and sex that could arise from differences in condition dependence of this trait between males and females. We found a positive relationship between UV chroma of breast feathers and hatching date. The amount of UV reflectance is thought to be negatively related to carotenoid content in feathers. The observed increase of UV chroma through the breeding season might therefore be caused by a seasonal decline in the availability and quality of Lepidoptera larvae – the main source of carotenoids in food of the Tits. We also observed a sex difference in the relationship between brightness of breast feathers (achromatic, structural component) and hatching date, which in males was negative and in females not significant. Our study provides further evidence that the timing of breeding is related to the expression of nestling carotenoid-based coloration, a potentially meaningful element of offspring–parent communication, and suggests a sex-specific effect of hatching date on its structural component.     

Achromatic Plumage Reflectance, Social Dominance and Female Mate Preference in Black-Capped Chickadees (Poecile atricapillus)

Individual variation in achromatic plumage reflectance of male Black‐capped Chickadees (Poecile atricapillus) is correlated with social rank and reproductive success, suggesting it may play an important role in sexual signaling. We asked whether female chickadees could assess male quality based on plumage, in the absence of information about relative social dominance. Sexually mature but inexperienced females captured during the pair formation period in late fall and early winter were presented with a choice of two unfamiliar, sexually experienced males in separate compartments of an outdoor mate choice arena. Following each preference trial, we released the males into a single compartment and scored their pairwise dominance interactions. In 10 of 11 trials, females spent significantly more time with the male subsequently identified as dominant, despite not witnessing the males interact. Spectral analysis of male plumage reflectance revealed that UV‐chroma of dark body regions (bib, cap and mantle) was significantly greater in dominant, preferred males and that dominant, preferred males had significantly brighter white cheek patches. There were no differences in vocalization rates of preferred and non‐preferred males. These results show that female chickadees can rapidly assess unfamiliar males based on visual cues, and suggest that variation in achromatic plumage functions in sexual signaling.     

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

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