Greater flamingos <Emphasis Type="Italic">Phoenicopterus roseus</Emphasis> use uropygial secretions as make-up

It was long thought that the colour of bird feathers does not change after plumage moult. However, there is increasing evidence that the colour of feathers may change due to abrasion, photochemical change and staining, either accidental or deliberate. The coloration of plumage due to deliberate staining, i.e. with cosmetic purposes, may help individuals to communicate their quality to conspecifics. The presence of carotenoids in preen oils has been previously only suggested, and here we confirm for the first time its presence in such oils. Moreover, the carotenoids in the uropygial secretions were the same specific pigments found in feathers. We show not only that the colour of feathers of greater flamingos Phoenicopterus roseus became more colourful due to the application of carotenoids from uropygial secretions over the plumage but also that the feathers became more colourful with the quantity of pigments applied over them, thus providing evidence of cosmetic coloration. Flamingos used uropygial secretions as cosmetic much more frequently during periods when they were displaying in groups than during the rest of the year, suggesting that the primary function of cosmetic coloration is mate choice. Individuals with more colourful plumage initiated nesting earlier. There was a correlation between plumage coloration before and after removal of uropygial secretions from feathers’ surfaces, suggesting that the use of these pigmented secretions may function as a signal amplifier by increasing the perceptibility of plumage colour, and hence of individual quality. As the cosmetic coloration strengthens signal intensity by reinforcing base-plumage colour, its use may help to the understanding of selection for signal efficacy by making interindividual differences more apparent.     

Relative importance of multiple plumage ornaments as status signals in golden whistlers (<Emphasis Type="Italic">Pachycephala pectoralis</Emphasis>)

Status signals are traits that advertise an individual’s competitive abilities to conspecifics during aggressive disputes. Most studies of status signals in birds have focussed on melanin-based plumage signals, but recent research shows that carotenoid-based signals may also play a role in aggressive signaling. We assessed the relative importance of melanin- and carotenoid-based plumage patches as agonistic signals in a small passerine, the golden whistler (Pachycephala pectoralis). Display signals in male golden whistlers include an unpigmented white throat patch, a carotenoid-based yellow breast and nape band, and a melanin-based black chin-stripe. We found that only the white throat patch was correlated with contest-related attributes. Males possessing large throat patches defended larger territories and commenced breeding earlier. When caged males with either experimentally reduced, or unmanipulated throat patches were presented to conspecifics, those with experimentally reduced patches attracted less aggression from male subjects. Focal males also responded faster to caged males with throat patches similar in size to their own, suggesting that they may assess relative throat patch size before engaging in aggressive encounters. Females did not discriminate between “reduced” or “control” treatments. Our data strongly suggest that only the unpigmented throat patch functions as a status signal. As this signal is unlikely to have significant development costs, honesty may be maintained through social costs.     

Delayed maturation of multiple signals in a migratory songbird

Many animals use multiple signals in sexual communication, but our understanding of the interactions between multiple signals, particularly in inexperienced breeders, is limited. In birds, delayed plumage maturation (DPM) is well documented; young birds appear duller than adults, despite reaching sexual maturity. Age-related changes in song structure are also common in songbirds, though the extent to which songs of yearling males differ from those of adults (delayed song maturation, DSM) and its prevalence in species with DPM is unknown. We tested for DSM in the mate-attraction (repeat) song of a species with dramatic DPM, the American redstart (Setophaga ruticilla). Repeat song structure of territorial yearling and adult males differed significantly, based on discriminant analysis of nine non-collinear song features. Combined with previous evidence of delayed maturation in territorial (serial) song, we provide the first evidence for DSM in different song types used in different behavioral contexts during the breeding season of a migratory songbird. Within adults, variation in repeat song was associated with pairing, earlier onset of breeding, and number of offspring sired, suggesting a potential benefit for expressing more adult-like song. We found no relationship between the expression of adult-like repeat song and plumage in either age class. These results indicate that delayed maturation of repeat song in yearling redstarts is not due to a generalized delay in signal development, and suggest further work is needed to determine whether mate attraction (for both yearling and adult males) provides greater pressure to sound more “adult-like” than to look more “adult-like”.     

Information content is more important than sensory system or physical distance in guiding the long-term evolutionary relationships between signaling modalities in Sceloporus lizards

Long-term signal evolution is shaped by a variety of selective pressures including the need to convey additional information or to improve message transfer to specific receivers or through multiple environments. Here, we test the relative importance of information and sensory modality in shaping the long-term evolution of multimodal signals in Sceloporus lizards. To broadcast identity at territorial boundaries, male Sceloporus use both visual motion (headbob) and chemical signals, whereas they use color (blue belly patches) to signal aggression. Using modern phylogenetic comparative methods, we found a negative correlation between evolutionary changes in visual motion (headbobs) and chemical (femoral pore) signals, but only indirect ties between the evolution of color and motion signals (both of which are perceived visually) through viviparity, and no evidence of an evolutionary link between color and chemical signals. We also find a negative correlation between arboreality and chemical signals. Thus, information content (in this case, broadcasting individual identity versus signaling aggression) appears to play a more important role than sensory modality or physical distance in guiding long-term signal evolution. Additional insights into the underlying evolutionary processes are described, illustrating the utility of a phylogenetic approach.     

Interacting effects of ambient light and plumage color patterns in displaying Wire-tailed Manakins (Aves,Pipridae)

Sending color signals to conspecifics may attract predators, leading to opposing selection pressures on the evolution of signal expression and display behavior in animals. The costs of signaling can be reduced, however, because conspicuousness is the combined result of the reflectance spectra of the displayer's color pattern and the spectra of ambient light illuminating the animal. Changes in ambient light can alter conspicuousness, even when chemical and structural color-generating mechanisms remain constant, potentially allowing animals to display their colors most fully in light environments where the benefits are greatest relative to the costs. Using spectroradiometric methods, we determined how light habitat use affects conspicuousness in adult males of the Wire-tailed Manakin Pipra filicauda, a lekking bird species with vivid plumage colors. We studied three aspects of visibility, including properties of the entire color pattern, visual contrast within an individual's plumage and a bird's contrast relative to its visual background. Wire-tailed Manakins usually displayed in forest shade environments, which reduced their conspicuousness at larger viewing distances, while maximizing visual contrast within the plumage color pattern at close viewing distances. Compared to sunspots, ambient light in forest shade reduces the contrast of individual bird colors with the background at close viewing distance. However, background contrast of individual bird colors in the shade was still relatively higher during sunny than during cloudy weather which may explain why males were more active when the sun was not blocked by clouds. Assuming that the visual perceptions of predators and other manakins do not differ from the reflectance patterns we measured, Wire-tailed manakins tend to display in light environments that reduce the conflicts between avoiding long distance detection by predators and displaying conspicuous color signals to visiting females.     

Do male plumage and song characteristics influence female off-territory forays and paternity in the hooded warbler?

Extra-pair fertilizations are common in many socially monogamous species, and paternity studies have indicated that females may use male vocal performance and plumage ornaments as cues to assess male quality. Female off-territory forays may represent a key component of female choice and male extra-pair mating success, and female foray behaviour is expected to be strongly influenced by indictors of male quality. In this study, we examined how male song and ornamentation affect how often females left their territories, which males they visited and extra-pair paternity in a socially monogamous passerine, the hooded warbler (Wilsonia citrina). We radiotracked 17 females during the fertile period and quantified male vocal performance (song output and rate) and plumage characteristics (size of the black melanin hood and colour of the black hood, yellow cheeks and breast areas). We obtained blood samples and determined paternity at 35 nests including those of 14 females that we radiotracked. Eleven (65%) of the 17 females forayed off-territory, whilst fertile and female foray rate was positively correlated with the number of extra-pair young in the nest. Females that left their territories more frequently were paired with males that sang at a low rate. In addition, extra-pair mates had higher song rates than the social mates they cuckolded (5.3 songs/min vs. 4.4 songs/min). Female off-territory forays or extra-pair paternity were not significantly related to male plumage characteristics. Our results indicate that a high song rate influences both the foray behaviour of a male’s social mate and the likelihood that he will sire extra-pair offspring with neighbouring females.     

Carotenoid-based status signalling in red-shouldered widowbirds (<Emphasis Type="Italic">Euplectes axillaris</Emphasis>): epaulet size and redness affect captive and territorial competition

In breeding plumage, the African male red-shouldered widowbirds (Euplectes axillaris) are black except for red carotenoid-based epaulets ('shoulder patches'), similar to the well-studied American red-winged blackbirds (Agelaius phoeniceus). To experimentally test the signal function of the red epaulets in male red-shouldered widowbirds, we manipulated epaulet colour and size (within natural variation), and observed the effects in two competitive contexts. First, in captivity, unfamiliar males with epaulets of different size or redness were staged in dyadic contests over food. Only epaulet manipulations significantly affected contest outcome; red epaulet males out-competed orange and control males, which in turn dominated blackened (no epaulet) males. Epaulet size manipulations had particularly strong effects; enlarged epaulet males vigorously defeated smaller epaulet males within colour treatments, whereas the independent effect of 'redness' (i.e., within size treatments) was much weaker. Second, in a field experiment, epaulets were manipulated prior to territory establishment. Males with enlarged epaulets acquired territories, whereas most males with blackened or reduced signals failed to obtain territories. Furthermore, among the territorial males, birds with enlarged epaulets defended larger territories and spent less time in boundary signalling, yet they received fewer intrusions from conspecifics. In contrast, epaulet manipulations had no effect on either female visitation rates or subsequent female settlement. These results concur with recent studies and suggest that the carotenoid-based epaulets of red-shouldered widowbirds are status signals, sexually selected through male contest competition for territories.     

The importance of white on black: unmelanized plumage proportion predicts display complexity in birds

Animal ritualized displays have been classically viewed as behavioral characters that decrease signal ambiguity or that facilitate the evaluation of costly exhibitions. It has been shown that their prevalence and level of complexity across species can reflect phylogenetic relationships between them, but the adaptive function of these behavioral traits is poorly known. Here, I hypothesize that, given that the efficacy of visual displays basically depends on conspicuousness and level of performance, species with low levels of conspicuousness may be forced to perform more complex varieties of a given display to get the same signal efficiency than other more conspicuous species. Thus, the evolution of display complexity, considered as the level of exaggeration of ritualized movements, may be explained as an adaptive trait and not only by phylogenetic inertia. I illustrate and test this hypothesis with the case of black-and-white plumage patches of pelecaniform birds. As predicted, there was a negative correlation between level of complexity and species conspicuousness (proportion of unmelanized plumage) for two different social displays. This indicates that classical ideas on the adaptiveness of ritualized displays should be considered to understand the present variation in signal form across species, which sheds light on the evolution of multiple signals.     

Strong effects of various incidence and observation angles on spectrometric assessment of plumage colouration in birds

The assessment of plumage colouration has become a common procedure in many ornithological studies, especially after the discovery of differences between avian and human visual systems, namely the avian ability to perceive ultraviolet wavelengths. There are many techniques available for plumage colour assessment. Nevertheless, unidirectional reflectance spectrometry seems actually to be the method of choice and is used in most studies. The active avian visual communication skills, the physical characteristics of the plumage surface and the birds’ body shape prompted us to study plumage colouration with different angles of incidence and observation by bidirectional reflectance spectrometry. In this study, we explore the effect of different angles of incidence and observation, on a single plane of incidence, on the reflectance spectrometric assessment of five differently coloured body regions of the blue fronted Amazon parrot (Amazona aestiva). Measurements were made using a AVS-USB2000 spectrometer with a DH-2000 light source and two different reflection probes (bifurcated fibre optic probe and a specially designed fibre holder). The results of this study illustrate clear angle dependence of colour parameters in differently coloured non-iridescent plumage regions of the blue fronted Amazon parrot. Our results may have direct implications on the observed transmitted information of display and its associated predation risk (trade-off) in birds, but also in other animals. Moreover, the results show that unidirectional reflectance spectrometry alone may be a rather limited procedure to quantify and compare true colour of a complex surface such as birds’ plumage. Therefore, slight but behavioural important characteristics of plumage pattern may remain undiscovered with this method.     

The effects of testosterone on antibody production and plumage coloration in male house sparrows (Passer domesticus)

Many bird species have patches of colour in their plumage, contrasting with their basic coloration, which are used to display and signal status to conspecifics. These are called ’badges of status’, because they are believed to be low-cost signals of social status. For a signalling system to be evolutionarily stable, cheating must be controlled. The conventional view is that there is frequent testing, which uncovers cheats. Recently, the immunocompetence handicap hypothesis (ICHH) suggested that signals may be dependent on testosterone for their development, with a cost being imposed through immune suppression. We report experiments on house sparrows (Passer domesticus) which show that testosterone significantly influences the size of the bib (a ’badge of status’). The ultimate effect of the testosterone manipulation was to impair antibody production, as predicted by the ICHH. However, testosterone manipulations also changed the levels of the ’stress hormone’ corticosterone. The level of corticosterone was also related to the degree of immunosuppression. After controlling for the effect of corticosterone, testosterone enhanced the birds’ ability to produce antibodies, counter to the ICHH. The hypothesis therefore must be modified. We suggest that testosterone has a dual effect: it leads to immunosuppression through a mechanism involving corticosterone but, conversely, leads to increased immunocompetence probably via dominance influencing access to resources. Received: 5 March 1999 / Received in revised form: 1 October 1999 / Accepted: 16 October 1999     

Ambient noise, motor fatigue, and serial redundancy in chaffinch song

Many animal signals are performed in a highly redundant manner as in some bird species where males sing several renditions of one song type before switching to another. However, differences in signal redundancy between contexts and between individuals are only poorly understood. We found that chaffinches (Fringilla coelebs) in noisier areas (i.e., close to waterfalls and torrents) sang longer bouts of the same song type before switching to a new type, suggesting that they use increased serial redundancy to get the message across in noisy conditions. This is the first evidence of a noise-dependent adjustment of signal redundancy in a songbird. In addition, we found that song types with faster trills were sung in shorter bouts suggesting that the performance of highly redundant song series is probably limited by motor constraints. Thus, in noisy environments, serial redundancy in bird song may reflect a trade-off between successful signal transmission and preventing motor fatigue.     

Ultraviolet reflectance influences female preference for colourful males in the European serin

Avian plumage colouration is one of the most impressive displays in nature and is frequently used as sexual signal. There is now considerable evidence that females consistently prefer males with the most elaborated colour displays. Bird colour vision expands into the ultraviolet (UV) range, which prompted several studies to test the importance of UV in mate choice, revealing that females are affected by the UV light component. These studies were mostly performed on structural plumage, whereas carotenoid-based plumage was rarely considered, although it also has a typical reflection peak in the UV. Our study tested the female choice over male yellow colouration, and whether it is influenced by UV removal, in the European serin (Serinus serinus), a sexually dichromatic cardueline finch, with males showing a conspicuous carotenoid-based yellow plumage. We shows that females preferred yellower males and that male attractiveness was lost when the UV colouration was blocked, with either of the UV-blocking techniques used. The results of our study indicate that the UV component of carotenoid colouration is important in the female mate assessment in serins and highlights the importance of considering colour perception in avian mate choice.     

Complex signal function: developing a framework of testable hypotheses

The basic building blocks of communication are signals, assembled in various sequences and combinations, and used in virtually all inter- and intra-specific interactions. While signal evolution has long been a focus of study, there has been a recent resurgence of interest and research in the complexity of animal displays. Much past research on signal evolution has focused on sensory specialists, or on single signals in isolation, but many animal displays involve complex signaling, or the combination of more than one signal or related component, often serially and overlapping, frequently across multiple sensory modalities. Here, we build a framework of functional hypotheses of complex signal evolution based on content-driven (ultimate) and efficacy-driven (proximate) selection pressures (sensu Guilford and Dawkins 1991). We point out key predictions for various hypotheses and discuss different approaches to uncovering complex signal function. We also differentiate a category of hypotheses based on inter-signal interactions. Throughout our review, we hope to make three points: (1) a complex signal is a functional unit upon which selection can act, (2) both content and efficacy-driven selection pressures must be considered when studying the evolution of complex signaling, and (3) individual signals or components do not necessarily contribute to complex signal function independently, but may interact in a functional way.Communicated by A. Cockburn     

Carotenoid-based colouration and ultraviolet reflectance of the sexual ornaments of grouse

Among the most familiar sexual signals are red, yellow, and orange sexual traits pigmented by carotenoids. Many birds can detect near-ultraviolet (UV) light, and UV signals can play key roles in mate choice. Grouse (Tetraonidae) exhibit bright carotenoid-dependent sexual ornaments, their supra-orbital combs, which to humans appear orange-red. Combs also reflect in the UV, which is not visible to humans but is likely to be visible to grouse. In male red grouse Lagopus lagopus scoticus, we show that comb UV reflectance decreases with increasing comb size and redness. By removing the epidermis of combs, where carotenoid pigments are, we show that the UV reflectance is a property of the dermis, underneath the red pigmented epidermis. Carotenoid pigmentation of combs acted as a mask to reduce reflectance by the dermis in the range 400–550 nm and in the UV, 300–400 nm. Patagium skin (non-ornamental skin under the wing) also reflects in the UV, but epidermis removal on this bare part tended to reduce UV reflectance, whereas removal of the red epidermis of combs increased UV reflectance. Males in better condition (greater body mass relative to size) had bigger and redder combs, but with less UV. Thus, carotenoid pigments of grouse combs are deposited on a white background with significant UV reflectance, which can influence how the signal is perceived by conspecifics. Carotenoid-based traits exhibit UV reflectance in a number of species, but how UV reflectance and carotenoid pigmentation influence colour remains little known for integumentary ornaments compared to plumage traits. UV vision is not uncommon in birds and other animals, so future studies should investigate how UV reflectance influences the perception of carotenoid-based signals of quality.     

Female yellowhammers (Emberiza citrinella) prefer yellower males: a laboratory experiment

The importance of male plumage coloration as a signal of male dominance and a cue for female choice in the monogamous yellowhammer, Emberiza citrinella, was tested in two sets of experiments in an indoor aviary. Dominance was tested by introducing two individuals, with no previous experience of each other, in an aviary with food and water. Aggression occurred more often between two males than between a male and a female and more frequently between pairs of males including at least one old male than between two young males. Dominance was not related to male colour in trials between males of the same age class, but it was in trials between an old and a young male, often differing in colour. Thus, age may be a more important determinant of dominance than colour. Female preference for more colourful males was tested by allowing hormone-induced females to choose between a more and a less colourful dummy male. Females spent more time in front of more colourful than drabber males and also more often perched beside colourful males than duller individuals. Although male colour cannot be ruled out as a dominance signal, the results suggest that male colour is primarily used as a signal in mate choice. Female choice may hence be responsible for maintenance of bright plumage in the male yellowhammer.     

Achromatic color variation in black-capped chickadees,Poecile atricapilla: black and white signals of sex and rank

Sexual dichromatism and phenotypic variation in elaborate male traits are common products of sexual selection. The spectral properties of carotenoid and structurally-based plumage colors and the patch sizes of melanin-based plumage colors have received considerable attention as sexual signals in birds. However, the importance of variation in achromatic plumage colors (white, gray and black) remains virtually unexplored, despite their widespread occurrence. We investigated a potential signal function of the achromatic black and white plumage of black-capped chickadees (Poecile atricapilla). We captured and color-banded 178 free-living chickadees and assessed winter flock dominance hierarchies by tabulating pairwise interactions at feeders. We recaptured 73 of these birds and measured plumage coloration for six body regions using a reflectance spectrometer and the area of melanin-based plumage patches from standardized photographs. We found extensive individual variation in chickadee plumage traits and considerable sexual dichromatism. Male black-capped chickadees have significantly brighter white plumage than females, larger black patches, and greater plumage contrast between adjacent white and black plumage regions. We also found rank differences in the plumage reflectance of males; high-ranking males, who are preferred by females as both social and extra-pair partners, exhibit significantly darker black plumage and grow their feathers more rapidly than low-ranking males. This variation among individuals reveals a potential signal function for achromatic plumage coloration in birds.     

Melanin-based colour polymorphism signals aggressive personality in nest and territory defence in the tawny owl (Strix aluco)

Nest and territory defence are risky and potentially dangerous behaviours. If the resolution of life history trade-offs differs between individuals, the level of defence may also vary among individuals. Because melanin-based colour traits can be associated with life history strategies, differently coloured individuals may display different nest and territory defence strategies. We investigated this issue in the colour polymorphic tawny owl (Strix aluco) for which plumage varies from dark to light reddish melanic. Accordingly, we found that (1) our presence induced a greater response (flying around) from dark-coloured than light-coloured females and (2) dark reddish males suffered lower nest predation rates than light-coloured males. In experimentally enlarged broods, the probability that females reacted after we played back the hoot calls of a stranger male was higher if these females were lighter reddish; the opposite pattern was found in experimentally reduced broods with dark parents being more reactive than light parents. Finally, darker females alarmed more frequently when paired with a light than with a dark male, suggesting that partners adjust their behaviour to each other. We also tested whether colouration is used as a signal by conspecifics to adjust the level of their defensive behaviour. Accordingly, breeding females responded more vigorously to a dark than a light reddish stuffed tawny owl placed beside their nest. We conclude that melanin-based colouration is a signal of alternative nest and territory defence behaviour that depends on ecological factors.     

Song amplitude and body size in birds

Bird song is a sexually selected multidimensional signal. A fundamental question regarding the evolution of sexually selected signals is what information they convey and how their honesty is maintained. Song amplitude is a performance-related signal trait that varies considerably between individuals, but this signal dimension has been neglected in past studies. I found that median song amplitude in male nightingales (Luscinia megarhynchos) and zebra finches (Taeniopygia guttata) did not vary significantly with body size or residual body mass. In contrast, I found a significant negative correlation between body size (and also residual mass) and the maximum song amplitude during interactive singing in nightingales. However, the function of these more subtle differences in song amplitude remains to be investigated. By and large, the results of this study suggest that mean song amplitude is unlikely to indicate a bird’s body size or current condition (measured as residual mass).     

Female brown-headed cowbirds' (<Emphasis Type="Italic">Molothrus ater</Emphasis>) social assortment changes in response to male song: a potential source of public information

In many species, females' behavior appears to be influenced by that of other females, particularly regarding mate choice. Females theoretically can reduce the costs associated with independent male assessment by observing conspecifics. Studies of brown-headed cowbirds suggest that females pay attention to other females' behavior. Group-housed females modify their song preferences, whereas females housed in pairs do not. What information is available to females in a group environment? To address this question, we studied two groups of juvenile (i.e. hatch-year birds) and adult female cowbirds in a naturalistic group setting. We used a longitudinal ABA design, consecutively introducing and removing males that differed in age, amount of song production and stage of song development, to isolate the male characteristics that related to changes in female behavior. Juvenile and adult females assorted by age class when singing adult males were in the aviary, but not when singing juveniles or silent males of any age class were in the aviary. Results from playback tests confirmed that adult male song alone influenced female age class assortment. Videotape analysis from playback tests revealed that females also wing stroked in response to male song. Other females sometimes approached females who wing stroked and observed them. We hypothesize that group-level changes in social organization and individual females' responses can serve as visual signals for other individuals.     

Parental effects on carotenoid-based plumage coloration in nestling great tits, Parus major

Carotenoid pigments have attracted much interest in behavioural and evolutionary ecology because of their dual function in immune physiology and as color signals. In vertebrates, carotenoids must ultimately be obtained from the diet, and the mechanisms and magnitude of this environmental dependence are central for understanding carotenoid signal functions and evolution. In the present cross-fostering experiment with great tits Parus major, we investigate pre- and postnatal parental effects (egg yolk carotenoids, parental coloration) on nestling size and carotenoid coloration, using HPLC analysis of egg yolk carotenoids, and a reflectance-based measure of ‘chroma’ that reflects the plumage pigment concentration. Both rearing environment and origin influenced offspring size and plumage chroma. Maternal allocation of carotenoids to eggs had a weak positive effect on nestling plumage chroma, whereas we found no prenatal maternal effects (egg size or yolk carotenoid concentration) on size. Nestling plumage chroma was also significantly predicted by the chroma of the rearing father, but not by the color of the rearing mother or either of the original (genetical) parents. Thus, both prenatal maternal effects and postnatal paternal effects influence the carotenoid-based plumage coloration of nestling great tits. Future studies will reveal if parental effects have long-term consequences for plumage development and associated fitness components.