Preen gland removal increases plumage bacterial load but not that of feather-degrading bacteria

The preen gland is a holocrine sebaceous gland of the avian integument which produces an oily secretion that is spread on the plumage during preening. It has been suggested that birds may defend themselves against feather-degrading bacteria (FDB) and other potential pathogens using preen gland secretions. However, besides some in vitro studies, the in vivo bacterial inhibitory effects of the preen oil on the abundance of feather-associated bacterial species has not yet been studied in passerines. Here we tested the effect of gland removal on the abundance of FDB and other-cultivable bacterial loads (OCB) of male house sparrows (Passer domesticus). Our results did not support earlier results on in vitro antibacterial activity of preen oil against FDB since the absence of the preen gland did not significantly affect their loads related to the control birds. In contrast, we found that preen gland removal led to higher loads of OCB. This result suggests that the antimicrobial spectrum of the preen oil is broader than previously thought and that, by reducing the overall feather bacterial loads, the preen gland could help birds to protect themselves against a variety of potentially harmful bacteria.     

Soil and preen waxes influence the expression of carotenoid-based plumage coloration

The signaling function of carotenoid-based plumage is mainly determined by the concentration of pigments in feathers. For this reason, most studies of the proximate control of coloration focus on processes during and preceding moult. In great tits Parus major, past research demonstrates that carotenoid-based plumage coloration honestly indicates male quality and, thus, may be a sexually selected signal. In this study, we investigate how dirt and preen oil influence the coloration of carotenoid-based feathers in the great tit. We collected six feathers from each individual bird; three feathers served as controls while the remaining three feathers were washed with a chloroform/methanol mixture to remove soil and preen waxes. We assessed plumage coloration using digital photography. This cleaning procedure slightly enhanced ornamentation; the experimentally cleaned feathers expressed hues shifted towards shorter wavelengths and expressed brighter overall coloration than control feathers. This is the first experimental study conducted on wild birds demonstrating that, in addition to pigment concentration, the presence of preen waxes and soils on feathers may contribute to variation in coloration.     

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.     

An individual and a sex odor signature in kittiwakes? Study of the semiochemical composition of preen secretion and preen down feathers

The importance of olfaction in birds’ social behavior has long been denied. Avian chemical signaling has thus been relatively unexplored. The black-legged kittiwake provides a particularly appropriate model for investigating this topic. Kittiwakes preferentially mate with genetically dissimilar individuals, but the cues used to assess genetic characteristics remain unknown. As in other vertebrates, their body odors may carry individual and sexual signatures thus potentially reliably signaling individual genetic makeup. Here, we test whether body odors in preen gland secretion and preen down feathers in kittiwakes may provide a sex and an individual signature. Using gas chromatography and mass spectrometry, we found that male and female odors differ quantitatively, suggesting that scent may be one of the multiple cues used by birds to discriminate between sexes. We further detected an individual signature in the volatile and nonvolatile fractions of preen secretion and preen down feathers. These results suggest that kittiwake body odor may function as a signal associated with mate recognition. It further suggests that preen odor might broadcast the genetic makeup of individuals, and could be used in mate choice to assess the genetic compatibility of potential mates.     

Hoopoes color their eggs with antimicrobial uropygial secretions

Uropygial gland secretions are used as cosmetics by some species of birds to color and enhance properties of feathers and teguments, which may signal individual quality. Uropygial secretions also reach eggshells during incubation and, therefore, may influence the coloration of birds’ eggs, a trait that has attracted the attention of evolutionary biologists for more than one century. The color of hoopoe eggs typically changes along incubation, from bluish-gray to greenish-brown. Here, we test experimentally the hypothesis that dark uropygial secretion of females is responsible for such drastic color change. Moreover, since uropygial secretion of hoopoes has antimicrobial properties, we also explore the association between color and antimicrobial activity of the uropygial secretion of females. We found that eggs stayed bluish-gray in nests where female access to the uropygial secretion was experimentally blocked. Furthermore, experimental eggs that were maintained in incubators and manually smeared with uropygial secretion experienced similar color changes that naturally incubated eggs did, while control eggs that were not in contact with the secretions did not experience such color changes. All these results strongly support the hypothesis that female hoopoes use their uropygial gland secretion to color the eggs. Moreover, saturation of the uropygial secretion was associated with antimicrobial activity against Bacillus licheniformis. Given the known antimicrobial potential of uropygial secretions of birds, this finding opens the possibility that in scenarios of sexual selection, hoopoes in particular and birds in general signal antimicrobial properties of their uropygial secretion by mean of changes in egg coloration along incubation.     

Assortative mating by carotenoid-based plumage colour: a quality indicator in American goldfinches,<Emphasis Type="Italic"> Carduelis tristis</Emphasis>

We investigated the mating patterns in 22 breeding pairs of wild American goldfinches (Carduelis tristis) with respect to their body size, condition, and carotenoid-based, yellow plumage colour. Using reflectance spectrometry, we objectively quantified plumage colours across the bird-visible wavelengths, revealing a unexpected UV peak in the reflectance spectrum from yellow feathers. We summarized our colour measurements using a principal components analysis to create a single variable, carotenoid PC1, that represents the intensity of this carotenoid-based yellow colour, a measure of phenotypic quality in this species. We found no evidence of assortative mating with respect to measures of body size or condition but there was positive assortative mating by carotenoid PC1, such that the yellow plumage colours of males and females were significantly correlated within pairs. We argue that the yellow carotenoid coloration of goldfinches may be important in mutual mate choice and, thus, that sexual selection in this species may act upon female ornamentation, as well as the more obvious plumage signals of males. Because assortative mating results in an increase in genetic variance, we suggest that this might be a mechanism that maintains variance in ornamental traits in spite of the variance-eroding effects of sexual selection.     

Comparative morphology and secretion chemistry of the scoli in caterpillars of<Emphasis Type="Italic"> Hyalophora cecropia</Emphasis>

The morphology of the variously coloured scoli (bristle-bearing structures on the integument, producing an exocrine secretion) on caterpillars of Hyalophora cecropia and the secondary chemistry of the discharged secretions have been investigated for the first time and compared. According to our scanning electron microscopic study, the red/orange, yellow and blue coloured groups of these glands differ morphologically. Gas chromatographic-mass spectrometric analyses showed that the patterns of secondary compounds in the respective glandular secretions are also different. Furthermore, the secretion of the penultimate larval instar is chemically distinct from that of the last instar, as are both secretions from the respective haemolymph. The results favour the idea that the differences in scoli colour, morphology and chemistry could affect various predator species differently.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

Beyond colour: consistent variation in near infrared and solar reflectivity in sunbirds (Nectariniidae)

The visible spectrum represents a fraction of the sun’s radiation, a large portion of which is within the near infrared (NIR). However, wavelengths outside of the visible spectrum that are reflected by coloured tissues have rarely been considered, despite their potential significance to thermal effects. Here, we report the reflectivity from 300 to 2100 nm of differently coloured feathers. We measured reflectivity across the UV-Vis-NIR spectra of different (a) body parts, (b) colour-producing mechanisms and (c) sexes for 252 individuals of 68 sunbird (family: Nectariniidae) species. Breast plumage was the most reflective and cap plumage the least. Female plumage had greater reflectivity than males. Carotenoid-based colours had the greatest reflectivity, followed by non-iridescent and iridescent melanin-based colours. As ordered arrays of melanin-filled organelles (melanosomes) produce iridescent colours, this suggests that nanostructuring may affect reflectance across the spectrum. Our results indicate that differently coloured feathers consistently vary in their thermal, as well as obvious visual, properties.     

Fifty shades of white: how white feather brightness differs among species

White colouration is a common and important component of animal visual signalling and camouflage, but how and why it varies across species is poorly understood. White is produced by wavelength-independent and diffuse scattering of light by the internal structures of materials, where the degree of brightness is related to the amount of light scattered. Here, we investigated the morphological basis of brightness differences among unpigmented pennaceous regions of white body feathers across 61 bird species. Using phylogenetically controlled comparisons of reflectance and morphometric measurements, we show that brighter white feathers had larger and internally more complex barbs than duller white feathers. Higher brightness was also associated with more closely packed barbs and barbules, thicker and longer barbules, and rounder and less hollow barbs. Larger species tended to have brighter white feathers than smaller species because they had thicker and more complex barbs, but aquatic species were not significantly brighter than terrestrial species. As similar light scattering principals affect the brightness of chromatic signals, not just white colours, these findings help broaden our general understanding of the mechanisms that affect plumage brightness. Future studies should examine how feather layering on a bird’s body contributes to differences between brightness of white plumage patches within and across species.     

Mimicry of queen Dufour's gland secretions by workers of Apis mellifera scutellata and A. m. capensis

The development of the Dufour's gland of workers of the two honey bee races Apis mellifera scutellata and A. m. capensis was measured. The Dufour's glands of A. m. capensis workers were longer and increased in length more rapidly than the glands of workers of A. m. scutellata at comparable ages. Analysis of the Dufour's gland secretions of workers and queens of both races revealed that there were caste and racial differences. Secretions of queenright A. m. scutellata workers were dominated by a series of long-chain hydrocarbons. In contrast the secretions of the A. m. capensis workers both under queenright and queenless conditions were a mixture of hydrocarbons and wax-type esters, as were those of queens. Multivariate analysis of the secretion profiles indicated that laying workers of both races mimic queens. The secretions of the A. m. capensis laying workers mimicked queen secretions most closely, enabling them to act as successful social parasites.     

Do spotless starlings place feathers at their nests by ultraviolet color?

A considerable number of bird species carry feathers to their nests. Feathers’ presence in the nests has traditionally been explained by their insulating properties. Recently, however, it has been suggested that feathers carried to the nests by females of the spotted starling (Sturnus unicolor L.) could have an ornamental function based on their ultraviolet (300–400 nm) and human-visible longer wavelength (400–700 nm) coloration. In our population, 95.7% of feathers found inside next-boxes occupied by nesting starlings were rock dove fly feathers. Of these feathers, 82.7% were naturally positioned with their reverse side oriented toward the entrance hole and 42.4% of all found feathers were situated within the nest-cup. Here we experimentally assess the signaling function of ultraviolet coloration of feathers in nests of spotless starlings by providing nests with a number of pigeon flight feathers that were respectively treated on their obverse, reverse, both, or neither side with a UV blocker. Starlings placed 42.5% of the experimental feathers in the nest-cup irrespective of the UV block treatment. Orientation of feathers toward the entrance hole was not related with their ultraviolet radiation. However, feathers placed within the nest-cup were more likely found with their reverse side oriented toward the entrance hole confirming our correlative findings. These results suggest a minor role of ultraviolet coloration on feather location by spotless starlings.     

Do feather-degrading bacteria actually degrade feather colour? No significant effects of plumage microbiome modifications on feather colouration in wild great tits

Parasites are known to exert selective pressures on host life history traits since the energy and nutrients needed to mount an immune response are no longer available to invest in other functions. Bird feathers harbour numerous microorganisms, some of which are able to degrade feather keratin (keratinolytic microorganisms) and affect feather integrity and colouration in vitro. Although named “feather-degrading” microorganisms, experimental evidence for their effects on feathers of free-living birds is still lacking. Here, we tested whether (i) keratinolytic microorganisms can degrade feathers in vivo and thus modify the colour of feathers during the nesting period and (ii) whether feather microorganisms have a long-term effect on the investment in colouration of newly moulted feathers. We designed treatments to either favour or inhibit bacterial growth, thus experimentally modifying plumage bacterial communities, in a wild breeding population of great tits (Parus major). Our analyses revealed no significant effects of the treatments on feather colours. Moreover, we found that differences in bacterial exposure during nesting did not significantly affect the colouration of newly moulted feathers. Our results suggest that significant feather degradation obtained during in vitro studies could have led to an overestimation of the potential of keratinolytic microorganisms to shape feather colouration in free-living birds.     

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.     

Alarm communication: a new function for the scent-gland secretion in harvestmen (Arachnida: Opiliones)

Most harvestmen are nocturnal, nonacoustical, and nonvisual arthropods. They have a pair of exocrine glands on the cephalothorax that produce defensive volatile secretions. We investigated in the field the possible alarm effect of these secretions in the gregarious harvestman Goniosoma aff. proximum. A cotton swab soaked with the species' own exudate (treatment), or with water (control), was held 1-2 cm from the center of harvestmen aggregations. The results showed that the gland secretion elicits an alarm response in Goniosoma: whereas 73.3% of the aggregations dispersed after being stimulated with the gland exudate, only 3.3% responded to the water control. Respondent groups are larger than nonrespondent groups, and the time of reaction to the secretion was inversely related to group size. This is the first demonstration of a chemically-mediated alarm effect in harvestmen. The alarm response in gregarious harvestmen has possibly evolved as a by-product of a primarily defensive reaction in the context of predator avoidance. The discovery of this novel function of scent-gland secretion is meaningful in view of the widespread occurrence of gregarious habit among species of the order Opiliones.     

Multiple UV reflectance peaks in the iridescent neck feathers of pigeons

Recent studies of colorful plumage signals in birds have been aided by the finding that birds can see ultraviolet (UV) light and thus may communicate using colors invisible to humans. Some of the pioneering and more pivotal work on avian color vision was performed with domestic pigeons (Columba livia), yet surprisingly there have been few detailed reports of the UV-reflecting properties of pigeon feathers. Here, I use UV-VIS fiber-optic spectrometry to document the full-spectrum reflectance characteristics of iridescent purple and green neck plumage in pigeons. Neck feathers that appear purple to the human eye exhibit four reflectance peaks—two in the UV and one in the blue and red regions—and thus exhibit a UV-purple hue. Neck feathers that appear green to the human eye are characterized by five spectral peaks: two in the UV (UVA and UVB), a predominant green peak, and secondary violet and red peaks, conferring a UV-purple-green color. Such elaborate UV coloration suggests that birds may use an even more complex and hidden UV signaling system than previously thought.     

A new maniraptoran dinosaur from China with long feathers on the metatarsus

The unusual presence of long pennaceous feathers on the feet of basal dromaeosaurid dinosaurs has recently been presented as strong evidence in support of the arboreal–gliding hypothesis for the origin of bird flight, but it could be a unique feature of dromaeosaurids and thus irrelevant to the theropod–bird transition. Here, we report a new eumaniraptoran theropod from China, with avian affinities, which also has long pennaceous feathers on its feet. This suggests that such morphology might represent a primitive adaptation close to the theropod–bird transition. The long metatarsus feathers are likely primitive for Eumaniraptora and might have played an important role in the origin of avian flight.Electronic Supplementary Material  Supplementary material is available for this article at     

The effect of metapleural gland secretion on the growth of a mutualistic bacterium on the cuticle of leaf-cutting ants

In Acromyrmex octospinosus leaf-cutting ants the metapleural glands produce an array of antibiotic compounds that serve as a general defence against unwanted microbes on the cuticle. Leaf-cutting ants also grow mutualistic Pseudonocardiaceae bacteria on their cuticle that produce antibiotics controlling a microfungal parasite of their fungus gardens. Interaction between this bacterium and gland secretion therefore seems unavoidable. We document the typical development of bacterial growth on the cuticle of young major workers, show that growth starts a few days after eclosion, and that the maximal cover is reached after 2–3 weeks and gradually declines when workers mature. Experimental closure of the metapleural glands had no effect on the initial exponential growth phase of the bacterium, but significantly reduced the cover during the decline phase. The age-dependent abundance of the bacterium and its partial dependence on metapleural gland secretion support the hypothesis that the abundance of this mutualist is actively regulated.     

Integrated plumage colour variation in relation to body condition, reproductive investment and laying date in the collared flycatcher

The possible integration of different sexual ornaments into a composite system, and especially the information content of such ornament complexes, is poorly investigated. Many bird species display complex plumage coloration, but whether this represents one integrated or several independent sexual traits can be unclear. Collared flycatchers (Ficedula albicollis) display melanised and depigmented plumage areas, and the spectral features (brightness and UV chroma) of these are correlated with each other across the plumage. In a 5-year dataset of male and female plumage reflectance, we examined some of the potential information content of integrated, plumage-level colour attributes by estimating their relationships to previous and current year body condition, laying date and clutch size. Females were in better condition the year before they became darker pigmented, and males in better current year condition were also darker pigmented. Female pigment-based brightness was positively, while male structurally based brightness was negatively related to current laying date. Finally, the overall UV chroma of white plumage areas in males was positively associated with current clutch size. Our results show that higher degree of pigmentation is related to better condition, while the structural colour component is associated with some aspects of reproductive investment. These results highlight the possibility that correlated aspects of a multiple plumage ornamentation system may reflect together some aspects of individual quality, thereby functioning as a composite signal.     

Skimming behaviour and spreading potential of Stenus species and Dianous coerulescens (Coleoptera: Staphylinidae)

Rove beetles of the genus Stenus Latreille and the genus Dianous Leach possess pygidial glands containing a multifunctional secretion of piperidine and pyridine-derived alkaloids as well as several terpenes. One important character of this secretion is the spreading potential of its different compounds, stenusine, norstenusine, 3-(2-methyl-1-butenyl)pyridine, cicindeloine, α-pinene, 1,8-cineole and 6-methyl-5-heptene-2-one. The individual secretion composition enables the beetles to skim rapidly and far over the water surface, even when just a small amount of secretion is emitted. Ethological investigations of several Stenus species revealed that the skimming ability, skimming velocity and the skimming behaviour differ between the Stenus species. These differences can be linked to varied habitat claims and secretion saving mechanisms. By means of tensiometer measurements using the pendant drop method, the spreading pressure of all secretion constituents as well as some naturally identical beetle secretions on the water surface could be established. The compound 3-(2-methyl-1-butenyl)pyridine excelled stenusine believed to date to be mainly responsible for skimming relating to its surface activity. The naturally identical secretions are not subject to synergistic effects of the single compounds concerning the spreading potential. Furthermore, evolutionary aspects of the Steninae’s pygidial gland secretion are discussed.