Optical properties of the uropygial gland secretion: no evidence for UV cosmetics in birds

Ultraviolet (UV) reflectance of the plumage is common in birds and plays an important role in sexual signalling. Recently, it has been proposed that birds are able to modify plumage UV reflectance by the application of uropygial gland secretion. Based on a survey of the optical properties of this secretion from 51 species belonging to 12 avian orders, we show that two main types of uropygial secretions exist, one predominantly found in passerines and one in non-passerines, both reducing relative UV reflectance of a white background (Teflon™ tape). We quantified how each type of secretion (exemplified by blue tit and mallard) affected feather UV reflectance. Both secretions reduced overall brightness and relative UV reflectance of white mallard feathers but hardly affected the reflectance of UV/blue blue tit crown feathers. According to models of avian colour vision, changes in reflectance due to application of the secretion were at or below the discrimination threshold of most birds. We conclude that the uropygial secretion is unlikely to play a major role in modifying plumage UV reflectance. However, the optical properties of the uropygial secretion may have been selected to interfere as little as possible with visual signaling through plumage reflectance. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Yolk testosterone organizes behavior and male plumage coloration in house sparrows (<Emphasis Type="Italic">Passer domesticus</Emphasis>)

Vertebrate eggs contain maternal steroid hormones in their yolks; and in avian species the concentration of these steroids vary within and among clutches. The organizational actions of these variable doses of maternal steroids in the avian egg, analogous to those regulating the development of sexually dimorphic traits, are little explored. In this study, we examined the organizational effects of yolk testosterone in the house sparrow, Passer domesticus, a sexually dichromatic passerine bird in which males are characterized by the presence of a throat patch or badge of black feathers that varies in size among males. We injected a physiological dose of testosterone or vehicle into eggs in the field, removed nestlings, and hand-raised them in the laboratory. At 5 months of age, we investigated treatment effects on plumage and behavior. Males that hatched from testosterone-injected eggs developed a larger badge than males that hatched from vehicle-injected (control) eggs. However, testosterone did not induce the expression of a badge in females. In staged dyadic encounters, both males and females hatched from testosterone-injected eggs were more successful at obtaining and defending a food source than individuals of the same sex hatched from control eggs. The results suggest that variable concentrations of maternal testosterone in the eggs of the house sparrow organize the expression of a plumage trait in males and behavior in both sexes.Communicated by J. Graves     

Age-dependent association between testosterone and crown UV coloration in male blue tits (Parus caeruleus)

The proximate basis of sexual traits can suggest mechanisms maintaining honesty in signalling. A central role hereby has been attributed to testosterone, although its importance for brightly coloured plumage has been questioned. We determined circulating testosterone levels in male blue tits captured at the start of breeding and demonstrated an age-dependent relationship between testosterone and male crown UV/blue coloration. In yearling males, testosterone increases with increasing ornamentation (higher UV chroma, higher chroma, more UV-shifted hue), whereas in older males, this relationship is negative, with less UV-ornamented males having higher testosterone. This pattern is robust since it occurred in 2 years, before and after egg laying, and in males sampled during the day and during the night, despite a tenfold difference in testosterone levels. Since more UV-ornamented young males gain higher within-pair paternity, while less UV-ornamented older males achieve more extra-pair matings, the results imply that higher testosterone is associated with reproductive success and attractiveness in both age classes. We hypothesise that this relationship could result from causal effects of testosterone on coloration or through associations with behaviour and suggest ways to test these hypotheses. Our results caution against premature dismissal of a potential role for testosterone in maintaining honesty of plumage signals.     

Extra-pair paternity and male characteristics in the pied flycatcher

The pied flycatcher (Ficedula hypoleuca) is sexually dichromatic with extreme variation in male plumage coloration. The benefit for males of having black plumage is controversial, and few studies have found evidence for a sexual selection benefit of being black rather than brown. However, blacker males may be better able to achieve extra-pair fertilizations (EPFs), which may be an important component of sexual selection. We studied the role of EPFs in sexual selection in the pied flycatcher by establishing a set-up where two males with different back coloration (blacker vs browner) bred simultaneously near each other. DNA fingerprinting analysis revealed that 11% of offspring resulted from EPFs, and that 22% of broods included extra-pair young (EPY) among 36 nests containing 223 nestlings. We found no evidence that browner males suffered more often from EPFs than blacker males. There was no correlation of male or female morphology or age with EPF frequency. However, breeding pairs with low genetic similarity had EPY in their nests significantly more often. Thus we argue that females paired with genetically dissimilar males may try to avoid the effects of extreme outbreeding by seeking extra-pair copulations (EPCs). Alternatively, incompatibility between genetically dissimilar mates may simply expose females to more extra-pair copulations.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

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.     

Carotenoids,immune response and the expression of sexual ornaments in male greenfinches (<Emphasis Type="Italic">Carduelis chloris</Emphasis>)

Allocation trade-offs of carotenoids between their use in the immune system and production of sexual ornaments have been suggested as a proximate mechanism maintaining honesty of sexual signals. To test this idea, we experimentally examined whether carotenoid availability in the diet was related to variation in antibody response to novel antigens in male greenfinches (Carduelis chloris aurantiiventris), a species with extensive carotenoid-dependent plumage colouration. We also measured the cost of mounting a humoral response in terms of circulating carotenoids. Finally, we examined the relationship between plumage colour, immune response and circulating carotenoids. We found that males with carotenoid-supplemented diets showed stronger antibody response than non-supplemented birds. We also found that activation of the immune system significantly reduced circulating carotenoids (24.9% lower in immune-challenged birds than in control birds). Finally, intensity (chroma) of ventral plumage colouration of males, a character directly related to concentration of total carotenoids in feathers, was negatively correlated with the immune response and circulating carotenoids in winter. These results support the idea that carotenoids are a limiting resource and that males trade ornamental colouration against immune response.     

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.     

An experimental test of female choice relative to male structural coloration in eastern bluebirds

Several experimental studies have shown that female birds use ornamental melanin and carotenoid plumage coloration as criteria in mate choice. Whether females choose mates based on natural variation in structural coloration, however, has not been well established. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet (UV)-blue plumage coloration on their head, back, wings, and tail, which is positively correlated with condition, reproductive effort, and reproductive success. We experimentally tested the hypothesis that female eastern bluebirds prefer as mates males that display brighter structural coloration by presenting breeding-condition females with males of variable coloration. We conducted two types of mate-choice experiments. First, females chose between males whose coloration was manipulated within the natural range of variation in the population; feathers were either brightened with violet marker or dulled with black marker. Second, females chose between males with naturally dull or bright plumage coloration. In both manipulated and unmanipulated coloration trials, female choice did not differ significantly from random with respect to structural coloration. We found no support for the hypothesis that the UV–blue coloration of male eastern bluebirds functions as a criterion in female mate choice.     

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.