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.     

Social environment during molt and the expression of melanin-based plumage pigmentation in male house sparrows (<Emphasis Type="Italic">Passer domesticus</Emphasis>)

Evolutionary biologists have shown much recent interest in the costliness and signal content of colorful plumage displays in birds. Although many studies suggest that both carotenoid- and structurally-based plumage colors are condition-dependent indicators of health and nutritional state at the time ornamental feathers are grown, there is little experimental evidence supporting the idea that melanin pigmentation is a reliable signal of condition during molt. Instead, melanin-based ornamental coloration often reveals the competitive ability and dominance of individuals throughout the year. However, this work does not indicate which proximate environmental factors shape the expression of melanin pigmentation at the time of feather growth. Because of the link between melanin coloration and the social environment, it is possible that the development of brightly colored plumage may be associated with aggressive social interactions during feather molt. Here, we show that melanin-based ornamental coloration in male house sparrows (Passer domesticus) is correlated with the degree to which individuals interact aggressively with conspecifics during molt. Males that were dominant (beta, but not alpha) within captive social groups during molt grew larger badges than subordinates. Groups of males that had higher rates of aggression during molt grew larger badges than less aggressive triads. To our knowledge, this is the first demonstration that melanin pigmentation and plumage-based status badges are related to the competitive history of individuals during feather development. By coupling badge size directly with aggressive experiences during molt, birds can use their status signal to honestly indicate their likelihood of winning agonistic encounters throughout the year.     

Foster,but not genetic,father plumage coloration has a temperature-dependent effect on offspring quality

Secondary sexual characters have most likely evolved through sexual selection because such traits indicate the genetic or phenotypic quality of the bearer. While genetic variation in such fitness-related traits should be depleted by directional selection, there are many cases in which variation is higher than expected. One hypothesis explaining this variation is that different phenotypes within a population are adapted to different environmental conditions. We tested this hypothesis by comparing the offspring quality of male pied flycatchers, Ficedula hypoleuca, with different degrees of melanin-based dorsal plumage coloration under different environmental conditions. To create different environmental growth conditions and to be able to separate offspring genotype effects from paternal effects on offspring body mass, we used a partial cross-foster design where the brood size was reduced or enlarged by one chick. We also examined the interactive effects of temperature and male phenotype because previous correlative studies suggest such temperature-dependent effects in this species. We show that, while manipulated brood size did not interact with male phenotype to affect offspring quality, temperature during the nestling period influenced the offspring quality of dark and brown foster (but not genetic) fathers. When the temperature was relatively low during the nestling period, foster offspring of black males were lighter than those raised by brown males; the opposite was true if temperature was relatively high. These results add a new aspect to our understanding of how variation in the degree of melanin-based coloration is maintained in wild populations and how phenotypic variation may be maintained in general.     

Eumelanin-based coloration and fitness parameters in birds: a meta-analysis

Although melanin is the most common pigment in animal integuments, the adaptive function of variation in melanin-based coloration remains poorly understood. The individual fitness returns associated with melanin pigments can be variable across species as these pigments can have physical and biological protective properties and genes involved in melanogenesis may vary in the intensity of pleiotropic effects. Moreover, dark and pale coloration can also enhance camouflage in alternative habitats and melanin-based coloration can be involved in social interactions. We investigated whether darker or paler individuals achieve a higher fitness in birds, a taxon wherein associations between melanin-based coloration and fitness parameters have been studied in a large number of species. A meta-analysis showed that the degree of melanin-based coloration was not significantly associated with laying date, clutch size, brood size, and survival across 26 species. Similar results were found when restricting the analyses to non-sexually dimorphic birds, colour polymorphic and monomorphic species, in passerines and non-passerines and in species for which inter-individual variation in melanism is due to colour intensity. However, eumelanic coloration was positively associated with clutch and brood size in sexually dimorphic species and those that vary in the size of black patches, respectively. Given that greater extent of melanin-based coloration was positively associated with reproductive parameters and survival in some species but negatively in other species, we conclude that in birds the sign and magnitude of selection exerted on melanin-based coloration is species- or trait-specific.     

Testosterone and melanin-based black plumage coloration: a comparative study

Despite the functional significance of melanin-based plumage coloration in social and sexual signaling, the mechanisms controlling its information content are poorly understood. The T-regulation hypothesis proposes that melanin ornaments signal competitive abilities via the effects of testosterone (T) mediating both melanization and sexual/aggressive behaviors. Using the phylogenetic comparative approach, we tested whether frontal black melanization is associated with elevated T around the time of breeding plumage development across all bird species with available T-data. We found a context-dependent relationship between melanization and T, varying with the type of ornamentation (patchy or full-black) and with the presumed taxonomic distribution of the hormonal control of plumage dichromatism. Within two taxa in which male plumage development is assumed androgen-dependent (Charadriiformes, Corvida), evolutionary increases in male melanization, and melanin dichromatism correlated with increases in T in most analyses but not within the basal lineage (ratites, Galloanseriformes) with androgen-independent male plumage. Among Passeroidea with presumably genetically or luteinizing-hormone-based male plumage, melanization and its dichromatism correlated with T only in species with <100% frontal melanization. These results were robust as we controlled for several confounding variables such as mating and parental behaviors. This study is the first to test and support the T-regulation hypothesis interspecifically, suggesting that among-species differences in melanization may have evolved in response to differences in circulating T in certain avian taxa. Our results imply that the extent of black ornamentation may serve as an honest indicator of male competitiveness in those species that evolved an appropriate hormonal basis (T dependence) for color production. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Carotenoid-derived ornaments reflect parental quality in male and female yellow-eyed penguins (<Emphasis Type="Italic">Megadyptes antipodes</Emphasis>)

The handicap principle suggests that ornamental traits that function as honest signals in mate selection must be costly to be effective. We evaluated in the sexually monochromatic yellow-eyed penguin (Megadyptes antipodes) whether the carotenoid-derived plumage and eye coloration predicts parental quality and whether males and females within pairs mate assortatively in relation to these carotenoid-derived ornaments. In addition, we investigated whether age or body condition was related to the coloration of the ornamental traits. In yellow-eyed penguins, parental quality of males and females was predicted by eye and head plumage coloration. Even when we controlled for gender- and age-specific differences, eye and head plumage coloration reflected honestly parental quality. Males and females mated assortatively in relation to these ornamental traits. While age influenced coloration of both the eye and head plumage, body condition was related only to the saturation of plumage coloration. These results provide evidence that the carotenoid-derived ornaments in yellow-eyed penguins reflect the parental abilities of birds and, therefore, may be costly signals. Potentially, female and male yellow-eyed penguins could use eye and plumage coloration as an indirect cue in assessing age and quality of individual birds during mate choice. This is only the second study to examine plumage coloration in relation to sexual selection in penguins, while conspicuous ornamental traits in other species of penguin beg the question whether they also play a role in sexual selection.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by C.R. Brown     

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.     

Elytra color as a signal of chemical defense in the Asian ladybird beetle Harmonia axyridis

Carotenoid- and melanin-based colors are valuable indicators of quality in many vertebrates, but their signaling role in invertebrates remains relatively unexplored. The Asian ladybird beetle Harmonia axyridis is an ideal organism for studies of this nature because males and females exhibit highly variable red and black colors on their elytra and are chemically defended with an alkaloid (harmonine). We used digital photography to quantify elytra coloration and absorbance spectrophotometry and gas chromatography mass spectrometry analyses to quantify pigment and alkaloid concentrations, respectively, in wild-caught male and female H. axyridis. We predicted that extensive or intense coloration would be an aposematic signal of high-alkaloid stores. We found that carotenoid pigments largely controlled variation in red elytra coloration. There was no relationship between alkaloid content and either elytra redness or carotenoid pigment concentration in either sex. However, we found a positive correlation between the extent of elytra coloration and alkaloid content. Animals with proportionally more red (or less black) on the elytra were more alkaloid-rich; this relationship was particularly strong in females. We also found that females with lighter black spots had greater amounts of harmonine than those with darker spots. These results suggest that elytra color patterns have the potential to reveal information about chemical defensiveness to mates or predators. Prior studies in this species show that nonmelanic forms are typically less active and yet more sexually attractive than melanic forms, and both results may be explained by the fact that nonmelanics are better chemically defended.     

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.     

Coloration in New World orioles

Summary Although sexual dichromatism in birds is usually ascribed to sexual selection, some workers argue that avian coloration is better explained by predation-related selection pressures. Supporting evidence for these latter hypotheses comes primarily from broad interspecific comparisons, which can be biased by a variety of factors. This study examines the predation-based hypotheses of Baker and Parker (1979), particularly the Predator Deflection Hypothesis, with reference to two closely-related oriole species: Icterus parisorum, which is dichromatic, and I. gularis, in which both sexes are brightly coloured. To test the prediction that bright coloration serves to divert the attention of predators away from cryptic young, rates of predation on nests of conspicuous (2 years of age or older) and dull-coloured (first-year). I. parisorum males were compared. The results showed equal predation on the young of males in both age/plumage classes. The Predator Deflection Hypothesis also predicts that, once a predator has been detected, brightly-coloured birds should attempt to distract it, whereas cryptic individuals should not. Tests using models of avian predators showed that this was not the case: cryptic I. parisorum females responded as aggressively toward the model as did conspicuous conspecific males. The same was true for I. gularis. Overall, the results did not support the idea that bright coloration has evolved in response to predation pressure in these species.     

Female siskins choose mates by the size of the yellow wing stripe

Commonly, female birds use the brightly coloured patches on males to choose the best-quality mates. Coloured wing patches, however, have received little attention or have been previously related to social behaviour (as a signal to recruit conspecific individuals at feeding patches) or foraging (to flush prey) contexts, rather than to sexual selection. Here we provide evidence that in siskins (Carduelis spinus), wing patches function in mate choice. Mate-choice experiments showed that females were attracted by the size of the yellow wing stripe of the male, but not by the size of its black bib, body size, general plumage brightness or age. Experiments on birds with manipulated yellow wing stripes showed that females were sensitive to the size of this colour patch, irrespective of other male qualities. The preference of female siskins for males with larger wing patches when searching for a mate may be explained by the relationship of this trait to foraging ability, which would ensure females good parental investment from the chosen male.Communicated by W.A. Searcy     

Feather coloration in museum specimens is related to feather corticosterone

Colorful ornaments in birds are often sexually selected signals of quality, and variation in ornament expression may be mediated by physiological stress through the secretion of corticosterone. However, testing for links between ornamentation and corticosterone often requires sampling live animals, and such physiological measures may not be matched in the time span in which they were sampled (e.g., very dynamic plasma corticosterone vs. plumage coloration, which is relatively static). Here, we use museum specimens to test for a link between the color of a sexual ornament and feather corticosterone at the time of ornament formation. In red-winged blackbirds, Agelaius phoeniceus, carotenoid-based epaulets appear to be important in male–male social interactions, territory maintenance, and female choice. We measured reflectance spectra of adult male epaulets and plucked adjacent feathers for corticosterone analysis via radioimmunoassay. We controlled for differences in the number of mates, specimen age, and geography by selecting only males with one mate and only birds collected in Florida during a 3-year period. Epaulet hue and red chroma did not vary with feather corticosterone, but males whose epaulets scored high for mean brightness and red brightness had significantly lower corticosterone than males with low brightness scores. This correlation with brightness but not hue or chroma is consistent with an effect of corticosterone (CORT) on feather microstructure, with elevated CORT leading to lower reflectance of white light from the keratin matrix surrounding the carotenoid pigments.     

Vivid birds respond more to acoustic signals of predators

Because conspicuous morphology such as colorful plumage may increase predation risk, we aimed to see if variation in plumage coloration could explain variation in avian anti-predator behavior. We included several measures of plumage coloration: human perception of vividness from images in field guides, total intensity from reflectance spectra of museum skins, contrasts calculated from physiological models of these spectra parameterized for both raptors and humans, chroma, and spectral saturation. We investigated how well these measurements predicted risk assessment in ten species of birds in St. John, U.S. Virgin Islands. We quantified how each species responded to playbacks of a predator’s calls and compared this response to that elicited by songs from a non-predatory, sympatric bird. We found that human-determined measures of vividness best predicted anti-predator responses of birds—more vividly colored species responded more to predators than duller species. No spectrophotometric variable explained variation in species reactions to a predator call. Our results suggest that vivid birds may compensate for their conspicuousness by being more responsive to the sound of predators and that more work is needed to better evaluate how animal coloration is quantified in comparative studies.     

The effects of preen oils and soiling on the UV–visible reflectance of carotenoid-pigmented feathers

Plumage coloration, particularly when carotenoid-based, is important in social signaling in birds. Although feather color is a relatively stable trait, individuals may modify it with “cosmetic” substances such as preen oils. In addition, dirt accumulation may influence plumage coloration and further affect signal perception by receivers. Here, we analyze the separate potential effects of preen oils and soil accumulation on the reflectance properties of carotenoid-pigmented feathers across the visual range of most bird species, which includes the ultraviolet (UV). Using the yellow portion of tail feathers of Bohemian waxwings (Bombycilla garrulus), we performed two separate experiments where: (a) preen oils and/or soil were removed, or (b) preen oils (from black-billed magpies Pica pica or eagle owls Bubo bubo) were added. Preen oil addition reduced brightness but increased UV hue and yellow chroma. UV chroma was reduced by the addition of magpie (but not owl) preen oil. Soil accumulation had little effect on plumage reflectance in the UV range but significantly reduced yellow chroma. According to models of avian vision, both of these effects are detectable by birds and biologically meaningful when compared with natural variation between the sexes and age classes. We conclude that preen oil and soil accumulation can significantly affect the UV–visible reflectance of carotenoid-based plumages. As such traits typically advertise individual quality, preening and soiling have the potential to modify the information content of carotenoid-based plumage traits and how these signals are perceived by receivers.     

Age before beauty? Relationships between fertilization success and age-dependent ornaments in barn swallows

When males become more ornamented and reproduce more successfully as they grow older, phenotypic correlations between ornament exaggeration and reproductive success can be confounded with age effects in cross-sectional studies, and thus say relatively little about sexual selection on these traits. This is exemplified here in a correlative study of male fertilization success in a large colony of American barn swallows (Hirundo rustica erythrogaster). Previous studies of this species have indicated that two sexually dimorphic traits, tail length and ventral plumage coloration, are positively correlated with male fertilization success, and a mechanism of sexual selection by female choice has been invoked. However, these studies did not control for potential age-related variation in trait expression. Here, we show that male fertilization success was positively correlated with male tail length but not with plumage coloration. We also show that 1-year-old males had shorter tails and lower fertilization success than older males. This age effect accounted for much of the covariance between tail length and fertilization success. Still, there was a positive relationship between tail length and fertilization success among older males. But as this group consisted of males from different age classes, an age effect may be hidden in this relationship as well. Our data also revealed a longitudinal increase in both tail length and fertilization success for individual males. We argue that age-dependent ornament expression and reproductive performance in males complicate inferences about female preferences and sexual selection.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Sexual selection and variation in reproductive strategy in male yellow warblers (Dendroica petechia)

Summary Male yellow warblers (Dendroica petechia) exhibit extensive variation in the amount and conspicuousness of the sexually distinctive brown streaking on the breast. We investigated this intraspecific variation in degree of sexual dichromatism to see if male plumage rank (essentially the amount of brown streaking) is correlated with the amount of reproductive effort allocated to parental investment, as sexual selection theory would predict. As predicted, the average rate of feeding visits to the nest by males was negatively correlated with the plumage rank of the male in 1982 (Fig. 1) and 1983 (Fig. 2), and varied little between years or among study areas. Within one study area, the higher nest visit rate of dull (low plumage rank) males was correlated with higher nestling growth rates. But in the other two study areas, where bright (high plumage rank) males predominated, nestlings of bright males tended to have the highest growth rates despite minimal nest visit rates. Thus, overall there was no correlation between male plumage rank and nestling growth rate, a potential index of relative reproductive success, in either year (Figs. 3 and 4). This overall equal nest success likely helps to maintain the behavioral and plumage polymorphism. Since we could find no evidence that the high nestling growth rates of bright males were due to an adjustment of female parental effort to compensate for low male parental effort, there must be some other benefit of being bright which contributes to nest success. To address this, we used the relationship between parental feeding rates and nestling growth rates for each nest as an index of relative territory quality. This analysis suggests that bright males generally occupy the best territories available, possibly because they are more aggressive in obtaining and defending them than dull males. Dull males appear to be relegated to poorer territories, where an increase in male parental contribution would be necessary to achieve high nestling growth rates. We propose that different males are using alternative but evolutionarily stable reproductive strategies, in a way that is consistent with the predictions of sexual selection theory. Allocation of limited reproductive effort to parental investment decreases as sexual dichromatism increases, which probably reflects an increased investment in intermale competition for the best territories.     

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

Background and temperature effects on Uca panacea color change

Organismal coloration is used for communication, camouflage, and thermoregulation. These functions of body coloration may impose conflicting demands upon color-changing organisms. Here, we examined interacting thermoregulatory and camouflage color change responses when fiddler crabs Uca panacea were subject to simultaneously changing temperatures (10, 25, 40 °C) and backgrounds (black, white). Crab coloration lightened on a white background and at high temperatures and darkened on a black background and at low temperatures, reflecting the camouflage and thermoregulatory functions of color change. Synergistic background and temperature treatments (i.e., hot/white or cold/black) induced strong color change responses. When temperature and background were in conflict (i.e., hot/black or cold/white), responses to background coloration constrained thermoregulatory color change such that carapace coloration did not change. Such conflicts are likely to be common in nature, especially in highly heterogeneous environments. Throughout the experiment, males remained lighter than females and showed a greater response to the hot/white treatment, driven by a stronger response to high temperatures. These differences may reflect the physiological, morphological, and behavioral differences associated with sexual selection and sexual dimorphism in this species.     

Quantifying iridescent coloration in animals: a method for improving repeatability

Quantification of animal colors is important to a variety of fields of research, especially those dealing with visual communication and sexual selection. Most animal colors are easily measured using well-established spectrophotometric techniques. However, the unique characteristics of iridescent colors present particular challenges and opportunities to quantify novel color metrics. Due to the fine-scale angle dependence of iridescent coloration, color metrics, such as hue and brightness, must be measured using methods that allow for repeatable comparison across individuals (e.g., by carefully controlling and measuring viewing geometry). Here, we explain how the optical characteristics of iridescent colors should be considered when developing measurement techniques, describe the pitfalls of some commonly used techniques, and recommend improved methods and metrics (angular degree of color change and breadth of reflectance) for quantifying iridescent color. In particular, most studies of iridescent birds to date have used less than ideal procedures and have not provided repeatability estimates for their methods. For example, we demonstrate here that measuring coloration from overlapping patches of iridescent feathers may be problematic, and we argue against methods that do not carefully control viewing geometry. We recommend measuring iridescence at maximal reflectance angles using an apparatus that allows for sample rotation, and we compare this technique to some other commonly used methods using iridescent gorget and crown feathers from Anna’s hummingbirds (Calypte anna). Our apparatus allows for the quantification of angular color change, and we found that maximal reflectance measurements using single feathers are highly repeatable both within feather samples and among samples within an individual.