Avian Colour Vision Research Articles

Avian nuptial plumage coloration changes differently when exposed to urban and rural areas

Avian plumage coloration could be altered by external factors like dust accumulation, but the effect of environmental conditions on such post-moult colour changes are poorly studied. Here, we investigated how exposure to the atmosphere in habitats of differing anthropogenic pressures modifies UV/blue structural plumage coloration — a sexually selected trait in eastern bluebirds Sialia sialis. We collected feather samples from live birds and distributed them in three types of habitats: (i) urban roads with low traffic, (ii) urban roads with high traffic, and (iii) rural roads with low traffic. After 4 weeks, the brightness of feathers decreased in all types of habitats-resulting in less-ornamented coloration. Changes in UV and blue chroma, however, varied with habitat type; chroma decreased in urban areas (becoming less ornamented), but increased in rural locations (becoming more ornamented). According to physiological models of avian colour vision, however, these changes in coloration tended to be below perceptual thresholds. Our results underscore the importance of how human activity can influence plumage colour of individual birds during the period between moult cycles which could have an impact on avian visual signalling. The study opens venues for further studies focusing on how the optical properties of specific airborne particles can influence coloration of wild-living animals.Significance statementPlumage coloration plays an important role in avian visual communication. In addition to moult-based change, coloration may change between moult cycles due to exposure to external factors like, for example, dust accumulation. Ecological factors modifying a passive plumage change, however, remains poorly studied. We collected samples of structurally coloured feather from a species wherein plumage colour is sexually selected and exposed them to outdoor conditions. We demonstrate that UV/blue chroma of coloration changes differently in urban and rural habitats. It increased in rural and decreased in urban areas. On the other hand, brightness decreased in both habitat types. Using physiological models of avian colour vision, however, we found that these colour changes were likely not perceptible to birds. Our study provides the first evidence that changes in coloration of fully grown feathers may be habitat dependent.

Red fruits exhibit lower colour diversity than red flowers as perceived by birds

Abstract Pollination and seed dispersal are crucial processes for plant reproduction, sharing ecological relevance and similarities, yet they have rarely been considered together. Flowers appear to express greater phenotypic diversity than fruits due to multiple confounding factors, which pose challenges for comparative analyses. The colours of flowers and fruits are important visual signal traits in pollination and seed dispersal, evolving under different selective pressures from their respective pollinators and seed dispersers. Birds constitute a unique plant‐interacting group that participates in both pollination and seed dispersal events. In this study, we focus on red flowers and red fruits associated with avian mutualists to gain insight into the intrinsic differences between flowers/pollination and fruits/seed dispersal. We conducted comparisons of colouration between 94 red flowers pollinated by birds and 99 red fruits dispersed by birds. The colour diversity was compared in both the spectral space and the avian colour vision spaces. Colour conspicuousness was analysed using avian colour vision models, as well as bee models. Pigeon colour preference was tested by controlled experiments utilizing red stimuli with and without secondary peaks at short wavelengths. Red fruits had lower colour diversity than red flowers, with redder hues and fewer secondary reflectance peaks. Avian colour vision models illustrated that fruits were more conspicuous than flowers achromatically, but not chromatically. Pigeons did not show preference for red with or without a secondary peak. Although both are red, there are significant differences between flowers and fruits in terms of colour diversity, spectral properties and colour perceptions. As we exclusively considered avian mutualists, these differences cannot be attributed to the differences in interacting animal groups or to their colour vision properties. This implies that the differences in evolutionary history between flowers and fruits may deserve further attention to understand the colour evolution. Read the free Plain Language Summary for this article on the Journal blog.

Carotenoid‐based plumage colour saturation increases with temperature in Australian passerines

AbstractAimBirds are often coloured yellow, olive, orange or red by carotenoids, which are plant pigments that are ingested with food. Here we test whether there are consistent patterns of intraspecific geographic variation in male carotenoid‐based plumage coloration linked to latitude, temperature, precipitation and primary productivity. It has been suggested that patterns of geographic variation should be different for yellow, compared to the metabolically derived red carotenoid‐based coloration, but this has not been tested.LocationAustralia.TaxaForty‐nine randomly selected species of Australian passerine birds (Passeriformes), 30 with yellow and 19 with red carotenoid‐based plumage coloration.MethodsCarotenoid‐based plumage patches of male museum specimens were measured using reflectance spectrometry, and spectra analysed using models of avian colour vision. For each species, we used linear models to determine whether geographic variation in carotenoid‐based colour correlated with latitude, temperature, precipitation or primary productivity. To determine whether effects are consistent across species, we used phylogenetic meta‐analyses.ResultsAlthough spatial environmental effects on carotenoid‐based plumage coloration varied across species, overall, species tended to have more saturated carotenoid‐based coloration at lower latitudes, and warmer regions. These effects applied mainly to resident species (as opposed to migratory or nomadic) and were stronger for red compared to yellow carotenoid‐based colours. We found no consistent effects of precipitation or primary productivity.Main conclusionsGeographic variation in male carotenoid‐based plumage coloration of Australian passerines is unlikely to be caused by variation in plant productivity (the ultimate source of carotenoids), and may be better explained by latitudinal gradients in the intensity of sexual selection or predation risk. Alternatively, temperature effects on red carotenoid‐based coloration may be driven by the hypothesized link between carotenoid and cellular metabolism. Latitudinal and temperature effects have the potential to generate perceivable differences in coloration across realistic spatial scales, contributing to divergence in visual signals.

Concerted Shifts in Absorption Maxima of Yellow and Red Plumage Carotenoids Support Specialized Tuning of Chromatic Signals to Different Visual Systems in Near-Passerine Birds

Recent evidence that absorption maxima (λRmin) expressed by colorful plumage pigments align to diagnostic cone sensitivities of affiliated visual systems suggests that birds employ specialized signals in relation to their color vision. However, these studies compared different pigments and clades for the violet (porphyrins in non-passerines) and ultraviolet (carotenoids in passerines) sensitive system, which confounds chemistry and phylogeny with tuning patterns. To test whether signal alignments to violet (VS) and ultraviolet (UVS) systems transcend confounding factors, parallel analyses were conducted for a diversity of near-passerines, a group in which plumage carotenoids occur in taxa with either visual system. Conventional and phylogenetically informed analyses confirmed earlier findings: short wavelength absorbing (yellow carotenoid) pigments aligned λRmin with the violet-sensitive (V) cone of VS species but with the short wavelength-sensitive (S) cone of UVS species, whereas long wavelength-absorbing (red carotenoid) pigments aligned only with the S cone of VS species. More extensive variation among VS yellow carotenoids produced λRmin alignments to cone sensitivities that differed at shorter (peaks) versus longer (overlaps) wavelengths. Ancestral trait reconstructions indicated that signals evolved to match pre-existing VS systems, but did not resolve scenarios for UVS systems. Regardless of historical details, alignments expressed a higher-level pattern in which λRmin values were blue-shifted for yellow and red carotenoids in VS compared to UVS species, a pattern opposite that expressed by receptor sensitivities between systems. Thus, generalized functional designs attributed to avian color vision allow for specialized visual communication through the development of chromatic signals suited to each perceptual system.

Carotenoid‐based plumage colouration is predicted by age and parasites in the male European serin

A fundamental assumption of theories on the evolution of sexual signals is that they should be costly to produce in order to honestly signal the quality of the sender. The expression of carotenoid‐based plumage signals is considered to be condition‐dependent, due to the role of carotenoids functioning as pigments and as health modulators. We assessed carotenoid‐based plumage colouration in relation to male condition in a free living population of male European serins Serinus serinus during the breeding season. Male serins were trapped for morphometric and colouration measurements, during a four‐year field study, in order to evaluate the signalling value of colouration in relation to body condition and parasites level. We compared two different forms of colour quantification based on spectral data – the most commonly used tristimulus colour variables and physiological models of avian colour vision – and found that they were highly correlated for this species. We investigated the signalling value of male plumage colouration and it was found to be related to age and ectoparasite load. Plumage double cone and patch size were negatively related to parasites level, whereas SWS ratio was positively related to parasites and age. Colouration was also related with the time since moult. Our results indicate that the colour expression of serin's plumage is age dependent and is related, in complex ways, with the ability to cope with parasitic infection.

Prey from the eyes of predators: Color discriminability of aposematic and mimetic butterflies from an avian visual perspective.

Predation exerts strong selection on mimetic butterfly wing color patterns, which also serve other functions such as sexual selection. Therefore, specific selection pressures may affect the sexes and signal components differentially. We tested three predictions about the evolution of mimetic resemblance by comparing wing coloration of aposematic butterflies and their Batesian mimics: (a) females gain greater mimetic advantage than males and therefore are better mimics, (b) due to intersexual genetic correlations, sexually monomorphic mimics are better mimics than female-limited mimics, and (c) mimetic resemblance is better on the dorsal wing surface that is visible to predators in flight. Using a physiological model of avian color vision, we quantified mimetic resemblance from predators' perspective, which showed that female butterflies were better mimics than males. Mimetic resemblance in female-limited mimics was comparable to that in sexually monomorphic mimics, suggesting that intersexual genetic correlations did not constrain adaptive response to selection for female-limited mimicry. Mimetic resemblance on the ventral wing surface was better than that on the dorsal wing surface, implying stronger natural and sexual selection on ventral and dorsal surfaces, respectively. These results suggest that mimetic resemblance in butterfly mimicry rings has evolved under various selective pressures acting in a sex- and wing surface-specific manner.

A vision physiological estimation of ultraviolet window marking visibility to birds.

Billions of birds are estimated to be killed in window collisions every year, worldwide. A popular solution to this problem may lie in marking the glass with ultraviolet reflective or absorbing patterns, which the birds, but not humans, would see. Elegant as this remedy may seem at first glance, few of its proponents have taken into consideration how stark the contrasts between ultraviolet and human visible light reflections or transmissions must be to be visible to a bird under natural conditions. Complicating matters is that diurnal birds differ strongly in how their photoreceptors absorb ultraviolet and to a lesser degree blue light. We have used a physiological model of avian colour vision to estimate the chromatic contrasts of ultraviolet markings against a natural scene reflected and transmitted by ordinary window glass. Ultraviolets markings may be clearly visible under a range of lighting conditions, but only to birds with a UVS type of ultraviolet vision, such as many passerines. To bird species with the common VS type of vision, ultraviolet markings should only be visible if they produce almost perfect ultraviolet contrasts and are viewed against a scene with low chromatic variation but high ultraviolet content.

The colorful language of Australian flowers

The enormous increase in phylogenetic information in recent years has allowed many old questions to be reexamined from a macroevolutionary perspective. We have recently considered evolutionary convergence in floral colors within pollination syndromes, using bird-pollinated species in Australia. We combined quantitative measurements of floral reflectance spectra, models of avian color vision, and a phylogenetic tree of 234 Australian species to show that bird-pollinated flowers as a group do not have colors that are significantly different from the colors of insect-pollinated flowers. However, about half the bird-pollinated flowers have convergently evolved a narrow range of colors with dominant long-wavelength reflection far more often than would be expected by chance. These convergent colors would be seen as distinctly different from other floral colors in our sample when viewed by honeyeaters (family Meliphagidae), birds with a phylogenetically ancestral type of color vision and the dominant avian pollinators in Australia. Our analysis shows how qualitative ideas in natural history, like the concept of pollination syndromes, can be given more precise definition and rigorous statistical testing that takes into account phylogenetic information.

ALL THE BETTER TO SEE YOU WITH: PIGMENT VARIATIONS IN PARROTS

Birds around the world rely on colourful plumage to identify and assess their kinsmen. Birds see colour using four colour visual pigments in their cone cells in addition to a rod pigment that is used in dim light vision. Each of these visual pigments is made up of one of five opsin proteins (LWS, SW1, SWS2, RH2 and RH1) and a chromophore, where each pigment absorbs a different range of light wavelengths. In some non-avian species the spectral sensitivities of these visual pigments have co-evolved with body colour and differ between individuals of the same species. Curiously, however, to date all studies suggest that the visual pigments vary little within the avian class, and even less within a particular species. To investigate further, Ben Knott, a researcher at Deakin University, and Wayne Davies, from the University of Western Australia, both in Australia, and their colleagues turned to the unusual parrot Platycercus elegans whose plumage can range from red to yellow (p. 4454).To begin with, the team characterised the wavelength sensitivity of the different visual pigments, by isolating rod and cone cells expressing the five different pigments. As expected, the five different pigments, absorbed light from the ultraviolet to the red ends of the visual light spectrum.Next, the team moved on to characterising the opsin proteins in more detail and sequenced the mRNA transcripts of the five opsins. As expected, three out of the five (LWS, SWS1 and SWS2) did not vary very much and were similar to opsins found in other bird species. However, the RH1 and RH2 opsins were noticeably different from the equivalent opsins in other birds, with proteins that would have extended carboxyl termini. What's more, the RH2 protein varied considerably, with mRNA transcripts encoding short, medium and long variants. The team suspect that by controlling the size of the RH2 opsin protein, parrots may be able to tweak the spectral sensitivity of the cone cell containing the RH2 visual pigment, thereby aiding them to see better in the yellow and red parts of the spectrum. Although the team still need to test this theory, it seems that there is more variation in avian colour vision than hitherto thought.

Color polymorphism in a land snail Cepaea nemoralis (Pulmonata: Helicidae) as viewed by potential avian predators.

Avian predation is one of the most probable factors maintaining polymorphism of shell coloration in Cepaea nemoralis. This assumption is justified by the fact that birds frequently forage on snails and their prey choice varies with morph coloration. However, in all preceding studies, the conspicuousness of morphs was determined only by using human vision which is significantly different from birds’ visual perception. In this study, we assessed how birds perceive colors of four Cepaea nemoralis morphs using physiological models of avian color vision. We calculated combined chromatic and achromatic contrast between shells and three habitat background types as a measure of shell conspicuousness. The degree of background color matching in Cepaea nemoralis depended on both shell morph and habitat type. On average, banded morphs were more conspicuous than unbanded morphs. Morphs were the most cryptic against dry vegetation and the most conspicuous on bare ground. We also found a significant interaction between habitat type and color morph. The relative conspicuousness of shell morphs depended on habitat and was the most variable against green vegetation. Our study provides the first insight into how potential avian predators view Cepaea nemoralis morphs. The results are discussed in light of multiple hypotheses explaining selective predation on Cepaea nemoralis morphs.

The phylogenetic distribution of ultraviolet sensitivity in birds

BackgroundColour vision in birds can be categorized into two classes, the ultraviolet (UVS) and violet sensitive (VS). Their phylogenetic distributions have traditionally been regarded as highly conserved. However, the complicated nature of acquiring spectral sensitivities from cone photoreceptors meant that until recently, only a few species had actually been studied. Whether birds are UVS or VS can nowadays be inferred from a wide range of species via genomic sequencing of the UV/violet SWS1 cone opsin gene.ResultsWe present genomic sequencing results of the SWS1 gene from 21 avian orders. Amino acid residues signifying UV sensitivity are found in the two most important spectral tuning sites 86 and 90 of Pteroclidiformes and Coraciiformes, in addition to the major clades, Palaeognathae, Charadriiformes, Trogoniformes, Psittaciformes and Passeriformes, where they where previously known to occur. We confirm that the presumed UVS-conferring amino acid combination F86, C90 and M93 is common to Palaeognathae and unique to this clade, despite available spectrometric evidence showing the ostrich retina to be VS.ConclusionsBy mapping our results together with data from previous studies on a molecular phylogeny we show that avian colour vision shifted between VS and UVS at least 14 times. Single nucleotide substitutions can explain all these shifts. The common ancestor of birds most likely had a VS phenotype. However, the ancestral state of the avian SWS1 opsin’s spectral tuning sites cannot be resolved, since the Palaeognathae are F86, C90 while the Neognathae are ancestrally S86, S90. The phylogenetic distribution of UVS and VS colour vision in birds is so complex that inferences of spectral sensitivities from closely related taxa should be used with caution.

Mimicry and masquerade from the avian visual perspective

Abstract Several of the most celebrated examples of visual mimicry, like mimetic eggs laid by avian brood parasites and palatable insects mimicking distasteful ones, involve signals directed at the eyes of birds. Despite this, studies of mimicry from the avian visual perspective have been rare, particularly with regard to defensive mimicry and masquerade. Defensive visual mimicry, which includes Batesian and Müllerian mimicry, occurs when organisms share a visual signal that functions to deter predators. Masquerade occurs when an organism mimics an inedible or uninteresting object, such as a leaf, stick, or pebble. In this paper, I present five case studies covering diverse examples of defensive mimicry and masquerade as seen by birds. The best-known cases of defensive visual mimicry typically come from insect prey, but birds themselves can exhibit defensive visual mimicry in an attempt to escape mobbing or dissuade avian predators. Using examples of defensive visual mimicry by both insects and birds, I show how quantitative models of avian color, luminance, and pattern vision can be used to enhance our understanding of mimicry in many systems and produce new hypotheses about the evolution and diversity of signals. Overall, I investigate examples of Batesian mimicry (1 and 2), Müllerian mimicry (3 and 4), and masquerade (5) as follows: 1) Polymorphic mimicry in African mocker swallowtail butterflies; 2) Cuckoos mimicking sparrowhawks; 3) Mimicry rings in Neotropical butterflies; 4) Plumage mimicry in toxic pitohuis; and 5) Dead leaf-mimicking butterflies and mantids.

Avian Vision and Egg Colouration: Concepts and Measurements

Avian egg colours and patterns are spectacularly diverse and provide a wonderful system to study both the functional and mechanistic basis of animal colouration. In order to do this, it is essential to consider the vision of the appropriate receiver, particularly birds, and to use methods and models to quantify the colour patterns independent of subjective human assessment. In this paper, I summarise some of the key features of avian colour, luminance, and pattern vision, and provide an outline of the methods available to quantify visual signals, including their associated benefits and limitations. In doing so, I make links to various recent studies of avian eggshell colouration, and a range of other relevant studies. Overall, I argue that considering receiver vision and employing appropriate methods to quantify eggshell appearance is crucial to understanding the basis of avian egg diversity.

How to evade a coevolving brood parasite: egg discrimination versus egg variability as host defences

Arms races between avian brood parasites and their hosts often result in parasitic mimicry of host eggs, to evade rejection. Once egg mimicry has evolved, host defences could escalate in two ways: (i) hosts could improve their level of egg discrimination; and (ii) negative frequency-dependent selection could generate increased variation in egg appearance (polymorphism) among individuals. Proficiency in one defence might reduce selection on the other, while a combination of the two should enable successful rejection of parasitic eggs. We compared three highly variable host species of the Afrotropical cuckoo finch Anomalospiza imberbis, using egg rejection experiments and modelling of avian colour and pattern vision. We show that each differed in their level of polymorphism, in the visual cues they used to reject foreign eggs, and in their degree of discrimination. The most polymorphic host had the crudest discrimination, whereas the least polymorphic was most discriminating. The third species, not currently parasitized, was intermediate for both defences. A model simulating parasitic laying and host rejection behaviour based on the field experiments showed that the two host strategies result in approximately the same fitness advantage to hosts. Thus, neither strategy is superior, but rather they reflect alternative potential evolutionary trajectories.

Evolution of ultraviolet vision in the largest avian radiation - the passerines

BackgroundInterspecific variation in avian colour vision falls into two discrete classes: violet sensitive (VS) and ultraviolet sensitive (UVS). They are characterised by the spectral sensitivity of the most shortwave sensitive of the four single cones, the SWS1, which is seemingly under direct control of as little as one amino acid substitution in the cone opsin protein. Changes in spectral sensitivity of the SWS1 are ecologically important, as they affect the abilities of birds to accurately assess potential mates, find food and minimise visibility of social signals to predators. Still, available data have indicated that shifts between classes are rare, with only four to five independent acquisitions of UV sensitivity in avian evolution.ResultsWe have classified a large sample of passeriform species as VS or UVS from genomic DNA and mapped the evolution of this character on a passerine phylogeny inferred from published molecular sequence data. Sequencing a small gene fragment has allowed us to trace the trait changing from one stable state to another through the radiation of the passeriform birds. Their ancestor is hypothesised to be UVS. In the subsequent radiation, colour vision changed between UVS and VS at least eight times.ConclusionsThe phylogenetic distribution of SWS1 cone opsin types in Passeriformes reveals a much higher degree of complexity in avian colour vision evolution than what was previously indicated from the limited data available. Clades with variation in the colour vision system are nested among clades with a seemingly stable VS or UVS state, providing a rare opportunity to understand how an ecologically important trait under simple genetic control may co-evolve with, and be stabilised by, associated traits in a character complex.

In the Eyes of the Beholders: Female Choice and Avian Predation Risk Associated with an Exaggerated Male Butterfly Color

Color ornaments are often viewed as products of countervailing sexual and natural selection, because more colorful, more attractive individuals may also be more conspicuous to predators. However, while evidence for such countervailing selection exists for vertebrate color ornaments (e.g., Trinidadian guppies), similar studies have yet to be reported in invertebrates. Indeed, evidence for female mate choice based on extant variation in male coloration is limited in invertebrates, and researchers have not explicitly asked whether more attractive males are also more conspicuous to predators. Here we provide evidence that more chromatic male cabbage white butterflies (Pieris rapae) are more attractive to females but should also be more conspicuous to predators. Female P. rapae preferentially mate with more chromatic males when choosing from populations of males with naturally occurring or commensurate, experimentally induced color variation. Mathematical models of female color vision confirm that females should be able to discriminate color differences between prospective mates. Further, chromatic and luminance contrast scores from female visual system models better predicted male mating success than did measures of male color derived more directly from color spectra. Last, models of avian color vision suggest that preferred males should be more conspicuous to known avian predators.

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

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

Visual modeling shows that avian host parents use multiple visual cues in rejecting parasitic eggs

One of the most striking outcomes of coevolution between species is egg mimicry by brood parasitic birds, resulting from rejection behavior by discriminating host parents. Yet, how exactly does a host detect a parasitic egg? Brood parasitism and egg rejection behavior provide a model system for exploring the relative importance of different visual cues used in a behavioral task. Although hosts are discriminating, we do not know exactly what cues they use, and to answer this it is crucial to account for the receiver's visual perception. Color, luminance ("perceived lightness") and pattern information have never been simultaneously quantified and experimentally tested through a bird's eye. The cuckoo finch Anomalospiza imberbis and its hosts show spectacular polymorphisms in egg appearance, providing a good opportunity for investigating visual discrimination owing to the large range of patterns and colors involved. Here we combine field experiments in Africa with modeling of avian color vision and pattern discrimination to identify the specific visual cues used by hosts in making rejection decisions. We found that disparity between host and foreign eggs in both color and several aspects of pattern (dispersion, principal marking size, and variability in marking size) were important predictors of rejection, especially color. These cues correspond exactly to the principal differences between host and parasitic eggs, showing that hosts use the most reliable available cues in making rejection decisions, and select for parasitic eggs that are increasingly mimetic in a range of visual attributes.

Can eggs in a cavity be a female secondary sexual signal? Male nest visits and modelling of egg visual discrimination in blue tits

Eggshell colouration is thought to function as a female-specific secondary sexual trait. While tests of this idea are rapidly accumulating in cavity-nesting birds, some fundamental underlying assumptions remain rarely investigated: namely, can males see eggshell coloration and perceive colour differences between the eggs of different females? We tested these two key assumptions in a natural population of blue tits (Cyanistes caeruleus). Using transponders, we tracked male nest visits and found that all males visited their nest-boxes while eggs were present and often visually accessible. Interestingly, some males also visited neighbouring nests. We then tested whether birds could detect eggshell coloration using models of avian colour vision; models were performed with and without limitations on visual performance owing to dim light. Both models found that differences in eggshell brightness were often easier to discriminate than differences in colour; there was more contrast in white eggshell background between clutches than within and its contrast against nest background was repeatable within clutches, suggesting these features could act as signals. Yet, the detectability of these contrasts depended entirely on model assumptions of visual limitations. Consequently, we need a better understanding of underlying visual mechanisms in dim-light environments and behavioural discrimination experiments before confirming the signalling potential of eggshell coloration.

Fat quill secretion in pigeons: could it function as a cosmetic?

Abstract In order to maintain functionality of the plumage, birds apply a variety of fatty substances to the feathers, usually derived from the preen gland. Such secretions could also have cosmetic functions, altering plumage reflectance, and there is indeed some evidence for such cosmetic functions of preen fats. Some birds, noticeably pigeons, use fats derived from modified feathers, so called fat quills, in a presumably similar fashion, but almost nothing is known about their putative functions. Here we present a first test of the possibility that fat quill secretion could function as a cosmetic. As models we used domestic pigeon races that produce fat quills, we first confirm their extensive use in plumage maintenance behaviour. We then assessed the effect of experimental addition and removal fat quill secretion on reflectance of white feathers using reflectance spectrometry and physiological models of avian colour vision. Addition of secretion to cleaned white feathers resulted in a significant, discriminable change in the feather reflectance spectrum, which was strikingly similar to the absorbance spectrum of passerine preen gland secretion. However, like previous studies on preen gland secretion, removal of fats from intact feathers did not significantly alter feather reflectance, indicating that fat quill secretion when present in quantities as applied by the birds does not noticeably affect plumage reflectance. Therefore we conclude that the potential for the fat quill secretion to modify plumage colour appears limited at best, and other functions, such as antibiotic defense against feather-degrading bacteria, should be considered.