Morphological variations in a widely distributed Eastern Asian passerine cannot be consistently explained by ecogeographic rules

Ecogeographic rules that describe quantitative relationships between morphologies and climate might help us predict how morphometrics of animals was shaped by local temperature or humidity. Although the ecogeographic rules had been widely tested in animals of Europe and North America, they had not been fully validated for species in regions that are less studied. Here, we investigate the morphometric variation of a widely distributed East Asian passerine, the vinous‐throated parrotbill (Sinosuthora webbiana), to test whether its morphological variation conforms to the prediction of Bergmann''s rule, Allen''s rules, and Gloger''s rule. We at first described the climatic niche of S. webbiana from occurrence records (n = 7838) and specimen records (n = 290). The results of analysis of covariance (ANCOVA) suggested that the plumage coloration of these parrotbills was darker in wetter/warmer environments following Gloger''s rule. However, their appendage size (culmen length, beak volume, tarsi length) was larger in colder environments, the opposite of the predictions of Allen''s rule. Similarly, their body size (wing length) was larger in warmer environments, the opposite of the predictions of Bergmann''s rule. Such disconformity to both Bergmann''s rule and Allen''s rule suggests that the evolution of morphological variations is likely governed by multiple selection forces rather than dominated by thermoregulation. Our results suggest that these ecogeographic rules should be validated prior to forecasting biological responses to climate change especially for species in less‐studied regions.     

A review of Gloger's rule,an ecogeographical rule of colour: definitions,interpretations and evidence

Gloger's rule is an ecogeographical rule that links animal colouration with climatic variation. This rule is named after C.W.L. Gloger who was one of the first to summarise the associations between climatic variation and animal colouration, noting in particular that birds and mammals seemed more pigmented in tropical regions. The term ‘Gloger's rule' was coined by B. Rensch in 1929 and included different patterns of variation from those described by Gloger. Rensch defined the rule in two ways: a simple version stating that endothermic animals are predicted to be darker in warmer and humid areas due to the increased deposition of melanin pigments; and a complex version that includes the differential effects of humidity and temperature on both main types of melanin pigments – eu‐ and phaeo‐melanin. The blackish eu‐melanins are predicted to increase with humidity, and decrease only at extreme low temperatures, while the brown‐yellowish phaeomelanins prevail in dry and warm regions and decrease rapidly with lower temperatures. A survey of the literature indicates that there is considerable variation/confusion in the way Gloger's rule is understood (based on 271 studies that define the rule). Whereas the complex version is hardly mentioned, only a quarter of the definitions are consistent with the simple version of Gloger's rule (darker where warm and wet), and most definitions mention only the effects of humidity (darker where wet). A smaller subset of studies define the rule based on other correlated climatic and environmental variables such as vegetation, latitude, altitude, solar radiation, etc., and a few even contradict the original definition (darker where cold). Based on the literature survey, I synthesised the qualitative (N = 124 studies) and quantitative (meta‐analytically, N = 38 studies, 241 effects) evidence testing the simple version of Gloger's rule (I found no tests of the complex version). Both lines of evidence supported the predicted effects of humidity (and closely linked variables) on colour variation, but not the effects of temperature. Moreover, humidity effects are not restricted to birds and mammals, as the data indicate that these effects also apply to insects. This suggests that the simple version of Gloger's rule as originally defined may not be valid, and possibly that the rule should be re‐formulated in terms of humidity effects only. I suggest, however, that more data are needed before such a reformulation, due to potential publication biases. In conclusion, I recommend that authors cite Rensch when referring to Gloger's rule and that they make clear which version they are referring to. Future research should concentrate on rigorously testing the validity and generality of both versions of Gloger's rule and establishing the mechanism(s) responsible for the patterns it describes. Since humidity seems to be the core climatic variable behind Gloger's rule, I suggest that the two most plausible mechanisms are camouflage and protection against parasites/pathogens, the latter possibly through pleiotropic effects on the immune system. Understanding the processes that lead to climatic effects on animal colouration may provide insights into past and future patterns of adaptation to climatic change.     

Reconciling ecogeographical rules: rainfall and temperature predict global colour variation in the largest bird radiation

Ecogeographical rules that associate climate with organismal form and function can reveal patterns of climatic adaptation. Two rules link animal coloration with climate: Gloger's rule (darker coloration where wet and warm), and Bogert's rule (darker coloration where cold). Whereas Gloger's rule was proposed for endotherms, and Bogert's rule for ectotherms, both rules may apply more broadly, despite their seemingly opposing effects. Here, we test this contradiction on a global scale across passerine birds. Consistent with Gloger's rule, birds were darker in wetter areas and, following Bogert's rule, lighter where warm, although birds became lighter again at very low temperatures. Rainfall and temperature had antagonistic or additive effects depending on their pattern of covariation, and this predicted whether birds followed the rules. We integrate both rules into a general framework to explain heterogeneity in climatic effects on coloration, which has implications to understand patterns of diversification, climatic adaptation and climate change impacts.     

Gloger's rule in plants: The species and ecosystem levels

Gloger''s rule posits that darker birds are found more often in humid environments than in arid ones, especially in the tropics. Accordingly, desert-inhabiting animals tend to be light-colored. This rule is also true for certain mammalian groups, including humans. Gloger''s rule is manifested at 2 levels: (1) at the species level (different populations of the same species have different pigmentation at different latitudes), and (2) at the species assembly level (different taxa at a certain geography have different pigmentation than other taxa found at different habitats or latitudes). Concerning plants, Gloger''s rule was first proposed to operate in many plant species growing in sand dunes, sandy shores and in deserts, because of being white, whitish, or silver colored, based on white trichomes, because of sand grains and clay particles glued to sticky glandular trichomes, or because of light-colored waxes. Recently, Gloger''s rule was shown to also be true at the intraspecific level in relation to protection of anthers from UV irradiation. While Gloger''s rule is true in certain plant taxa and ecologies, there are others where “anti-Gloger” coloration patterns exist. In some of these the selective agents are known and in others they are not. I present both Gloger and “anti-Gloger” cases and argue that this largely neglected aspect of plant biology deserves much more research attention.     

Male achromatic wing colouration is related to body condition and female reproductive investment in a dichromatic species,the upland goose

In many bird species, achromatic plumage patch size can serve as a male status signal, but the use of variations in the achromatic colours themselves as a quality signal has only recently come into focus. In our study, we sought to determine whether achromatic plumage reflects individual quality in the upland goose (Chloephaga picta leucoptera). We examined the relationship between male head and wing reflectance, male condition and female reproductive investment. We found that males with darker specula and greater contrast between the white wing coverts and the speculum were in a better body condition. Variations in the brightness of the white plumage were not a quality signal in the upland goose. The information gleaned from the wing colouration of male upland geese could be used during mate selection, when females choose their mate on the basis of the outcomes of aggressive encounters. During these fights, the males expose their white coverts and their specula, which are normally tucked beneath body feathers.     

Achromatic plumage reflectance predicts reproductive success in male black-capped chickadees

The size of achromatic (black, white, gray) plumage patchesserves as a male status signal in many species of birds, butvariation in the colors of these patches has received littleattention. We assessed the relation between achromatic plumagereflectance, dominance rank, body condition, and reproductivesuccess in male black-capped chickadees, Poecile atricapillus.We measured plumage reflectance for five body regions of 40male chickadees in late winter and monitored these males throughoutthe following breeding season to determine whether they survivedto breed, whether they successfully paired, whether their partnerlaid eggs, and both their apparent and realized reproductivesuccess. As expected from past studies, a male's dominance ranksignificantly predicted whether his partner laid eggs. However,only achromatic plumage reflectance significantly predictedother measures of male reproductive performance. Among maleswho fledged at least one offspring, both the brightness of whiteplumage regions and the UV-chroma of melanin-based plumage regionswere significant predictors of the proportion of within-pairyoung in their nests. When we consider all males we measured,assigning zero values to males who failed to sire any offspring,the UV-chroma of melanin-based plumage regions was a significantpredictor of realized reproductive success. Bib size was alsorelated to male realized reproductive success. Our findingssuggest that individual variation in achromatic plumage mayplay an important role in sexual signaling in chickadees.     

Adaptation of Pelage Color and Pigment Variations in Israeli Subterranean Blind Mole Rats,Spalax Ehrenbergi

Background

Concealing coloration in rodents is well established. However, only a few studies examined how soil color, pelage color, hair-melanin content, and genetics (i.e., the causal chain) synergize to configure it. This study investigates the causal chain of dorsal coloration in Israeli subterranean blind mole rats, Spalax ehrenbergi.

Methods

We examined pelage coloration of 128 adult animals from 11 populations belonging to four species of Spalax ehrenbergi superspecies (Spalax galili, Spalax golani, Spalax carmeli, and Spalax judaei) and the corresponding coloration of soil samples from the collection sites using a digital colorimeter. Additionally, we quantified hair-melanin contents of 67 animals using HPLC and sequenced the MC1R gene in 68 individuals from all four mole rat species.

Results

Due to high variability of soil colors, the correlation between soil and pelage color coordinates was weak and significant only between soil hue and pelage lightness. Multiple stepwise forward regression revealed that soil lightness was significantly associated with all pelage color variables. Pelage color lightness among the four species increased with the higher southward aridity in accordance to Gloger''s rule (darker in humid habitats and lighter in arid habitats). Darker and lighter pelage colors are associated with darker basalt and terra rossa, and lighter rendzina soils, respectively. Despite soil lightness varying significantly, pelage lightness and eumelanin converged among populations living in similar soil types. Partial sequencing of the MC1R gene identified three allelic variants, two of which were predominant in northern species (S. galili and S. golani), and the third was exclusive to southern species (S. carmeli and S. judaei), which might have caused the differences found in pheomelanin/eumelanin ratio.

Conclusion/Significance

Darker dorsal pelage in darker basalt and terra rossa soils in the north and lighter pelage in rendzina and loess soils in the south reflect the combined results of crypsis and thermoregulatory function following Gloger''s rule.     

The dark side of coloration: Ecogeographical evidence supports Gloger's rule in American marsupials

Geographical distribution of color phenotypes and associations with ecological predictors remains poorly understood. An important geographic pattern concerning this topic is Gloger's rule, which predicts the increase of pigmentation in endothermic animals from cold and dry to warm and wet environments. Didelphid marsupials exhibit a variety of color patterns, ranging from light and dark uniform to more complex colorations. However, surprisingly little is known about the adaptive significance of dark coloration in this singular group of mammals. Using a phylogenetic comparative approach, we investigated whether coloration in different body regions of didelphids (i.e., dorsum and face) is associated with variables representing heat and humidity of the environment, as predicted by Gloger's rule. We demonstrated that Gloger's rule explains the interspecific color variation in American marsupials, especially when considering the facial region. Thus, dark coloration was more frequent among didelphid species occupying warm and wet environments than cold and dry environments. We also discuss the selective forces that can potentially explain coat color variation in didelphid marsupials, including camouflage, pathogen resistance, and pleiotropy hypotheses.     

Why do melanin ornaments signal individual quality? Insights from metal element analysis of barn owl feathers

Melanin-based variation in colour patterns is under strong genetic control and not, or weakly, sensitive to the environment and body condition. Current signalling theory predicts that such traits may not signal honestly phenotypic quality because their production does not entail a significant fitness cost. However, recent studies revealed that in several bird species melanin-based traits covary with phenotypic attributes. In a first move to understand whether such covariations have a physiological basis, we quantified concentrations of five chemical elements in two pigmented plumage traits in the barn owl (Tyto alba). This bird shows continuous variation from immaculate to heavily marked with black spots (plumage spottiness) and from dark reddish-brown to white (plumage coloration), two traits that signal various aspects of individual quality. These two traits are sexually dimorphic with females being spottier and darker coloured than males. We found an enhancement in calcium and zinc concentration within black spots compared with the unspotted feather parts. The degree to which birds were spotted was positively correlated with calcium concentration within spots, whereas the unspotted feather parts of darker reddish-brown birds were more concentrated in zinc. This suggests that two different pigments are responsible for plumage spottiness and plumage coloration. We discuss the implications of our results in light of recent experimental field studies showing that female spottiness signals offspring humoral response towards an artificially administrated antigen, parasite resistance and fluctuating asymmetry of wing feathers.An erratum to this article can be found at     

THE BIRDS OF TASMANIA: ECOLOGY AND EVOLUTION

SUMMARY Tasmania, about the size of Ireland, separated from the Australian mainland by 140 miles of sea, and isolated for about the last 12,000 years, has 104 species of native breeding land-birds, with the addition of ten introduced species. The environment is described; in particular the vegetation is classified into nine natural types and three produced by European man; and the distribution of the bird species among these is defined and discussed. The other vertebrates are briefly considered. No extinction is definitely known to have taken place as a result of European settlement except of the Tasmanian Emu (and of course Tasmanian Man). An attempt is made to reconstruct the Late Pleistocene history of Tasmania and its vegetation, with special reference to the Last Glaciation, when the island would have been joined to the mainland. When the avifauna is divided into categories, water-birds, raptors, etc., it is found to have much the same proportional composition as the Australian mainland avifaunas with which it is compared, though it consists of many fewer species. The vegetation types of the colder and wetter areas of Tasmania house far fewer species of birds than the drier and warmer habitats. The 104 breeding species include 14 endemics, which are considered in detail, and 27 endemic subspecies. As shown by comparison with other islands, the total proportionate endemism is extraordinarily high for a recent continental island (though it is actually lower than that in the remote ecological island formed by the sclerophyll of southwestern Western Australia). A contributory cause may be that Tasmania is not regularly visited by land-birds from the continent (though at least one-fifth of the Tasmanian species are partial or total migrants in winter). The most noteworthy endemics are two monotypic genera, Lathamus and Acanthornis, and the Native Hen Tribonyx mortierii, which has become flightless apparently in the face of a formidable array of local predators. Considerations of climate and habitat suggest that at least half the avifauna, including 19 of the endemic subspecies and six of the members of superspecies, arrived in Tasmania some time after the amelioration of the Last Glaciation began, some 18,000 years ago. Geographical considerations suggest that four of those six members of superspecies existed in their present form when the land-bridge to the mainland was cut, 12,000 years ago. Certain habitats, widespread over southeastern Australia during the glaciation, are now virtually confined to Tasmania where they form the stronghold of certain species, such as the Pink Robin Petroica rodinogaster, which occur only as relicts on the mainland. The endemic Scrub Tit Acanthornis magnus is practically confined to such habitats, where its ecology suggests it would have been well adapted to glacial conditions. Among the local endemics there is a strong tendency for colouration to be more saturated than on the continent of Australia (Gloger's rule) but there is less consistency in tendency to greater size (Bergmann's rule). A noteworthy proportion of the Tasmanian species have duller plumage than their mainland relatives, usually with an assimilation to female or juvenile plumage—unexpected in an island as big as Tasmania with so considerable an avifauna.     

Revisiting the condition‐dependence of melanin‐based plumage

Support against the condition‐dependence of melanin plumage signals has relied on data from species exhibiting both melanin‐ and carotenoid‐based plumage ornaments. As the mechanisms leading to variation in carotenoid‐ and melanin‐based plumage differ fundamentally, these systems may not be ideal to assess the condition‐dependence of melanin signals. Instead, we hypothesized that melanin‐plumage is more likely to signal condition in purely achromatic species. We performed a meta‐analysis reviewing evidence for condition‐dependent melanic plumage: we compared the net effect size for the relationship between melanin traits and condition in species that are achromatic versus species that also display a carotenoid‐based trait. Our results indicate that melanin plumage is condition‐dependent in species of both plumage types. Contrarily to our prediction, this finding suggests that melanin ornament condition‐dependence is not conditional on the context of other ornaments within a species. Instead, melanin ornaments should be viewed as potential condition‐dependent signals in all species.     

Frequency distribution and environmental correlates of plumage polymorphism in the grey fantail Rhipidura fuliginosa

Abstract

The grey fantail (Rhipidura fuliginosa) in New Zealand displays a striking plumage polymorphism. Some individuals are coloured almost entirely black (the “black morph"), while other individuals sport a contrasting brown and white plumage (the “pied morph"). The adaptive significance of plumage polymorphism in this species is unknown. We mapped the relative distribution and frequency of each morph across the entire South Island range of the fantail, and correlated the frequency of the morphs with a variety of ecological variables. The black morph comprised <5% of individuals across the South Island and, contrary to previous observations, was least frequent at the southern extremes of its range. From historical records, the frequency of the black morph also appears to have declined, although we cannot rule out a bias in reporting rates of the black morph in the literature. The relative frequency of the two morphs was not related to vegetation type, annual rainfall, altitude, or mean annual temperature. Although we could not identify an environmental variable that might explain the distribution of the two morphs over the South Island, changes in the relative abundance of each morph suggest a dynamic process that warrants further long‐term study.     

Climatological correlates for body size of five species of Australian mammals

Size variation of body and skull of five species of Australian mammals (echidna, Tachyglossus aculeatus ; brush-tail possum, Trichosurus vulpecula ; eastern grey kangaroo, Macropus giganteus ; western grey kangaroo, M. fuliginosus ; red kangaroo, M. rufus ), is related to climatic factors. All five species show trends in body size that conform with Bergmann's rule, individuals from colder environments being larger than those from warmer areas. The western and eastern grey kangaroos also conform with Allen's rule, the relative size of their extremities being large in warmer areas. In four of the five species (not the red kangaroo) body size is also correlated with indices of biomass productivity. However, since biomass productivity and ambient temperature are related to some extent, it is difficult to separate the effects of these factors.     

Macro‐ and microclimatic interactions can drive variation in species' habitat associations

Many species are more restricted in their habitat associations at the leading edges of their range margins, but some species have broadened their habitat associations in these regions during recent climate change. We examine the effects of multiple, interacting climatic variables on spatial and temporal patterns of species' habitat associations, using the speckled wood butterfly, Pararge aegeria, in Britain, as our model taxon. Our analyses reveal that this species, traditionally regarded as a woodland‐dependent insect, is less restricted to woodland in regions with warmer winters and warmer and wetter summers. In addition, over the past 40 years of climate change, the species has become less restricted to woodland in locations where temperature and summer rainfall have increased most. We show that these patterns arise mechanistically because larval growth rates are slower in open (i.e. nonwoodland) habitats associated with colder microclimates in winter and greater host plant desiccation in summer. We conclude that macro‐ and microclimatic interactions drive variation in species' habitat associations, which for our study species resulted predominantly in a widening of habitat associations under climate change. However, species vary in their climatic and nonclimatic requirements, and so complex spatial and temporal patterns of changes in habitat associations are likely to be observed in future as the climate changes.     

Spatial patterns in the size of Chinese lizards are driven by multiple factors

BackgroundFor almost two centuries, ecologists have examined geographical patterns in the evolution of body size and the associated determinants. During that time, one of the most common patterns to have emerged is the increase in body size with increasing latitude (referred to as Bergmann''s rule). Typically, this pattern is explained in terms of an evolutionary response that serves to minimize heat loss in colder climates, mostly in endotherms. In contrast, however, this rule rarely explains geographical patterns in the evolution of body size among ectotherms (e.g., reptiles).LocationChina.AimIn this study, we assembled a dataset comprising the maximum sizes of 211 lizard species in China and examined the geographical patterns in body size evolution and its determinants. Specifically, we assessed the relationship between body size and climate among all lizard species and within four major groups at both assemblage and interspecific levels.ResultsAlthough we found that the body size of Chinese lizards was larger in warmer regions, we established that at the assemblage level, size was correlated with multiple climatic factors, and that body size–climate correlations differed within the four major groups. Phylogenetic analysis at the species level revealed that no single climatic factor was associated with body size, with the exception of agamids, for which size was found to be positively correlated with temperature.Main conclusionsGeographical patterns in Chinese lizard body size are driven by multiple factors, and overall patterns are probably influenced by those of the major groups. We suggest that our analyses at two different levels may have contributed to the inconsistent results obtained in this study. Further studies investigating the effects of altitude and ecological factors are needed to gain a more comprehensive understanding of the evolution of ectotherm body size.     

Latitudinal variation of morphological characteristics in the swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae)

Patterns of latitudinal variation in the phenotype or genotype of an organism may provide evidence for natural selection. In this study, we investigated seven populations of swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae), a non‐migratory species, to explore the latitudinal variation of morphological characteristics in adults. The results showed that body size and the development of dark pigmentation on wings in this species responded strongly to latitude. The body size of both male and female adult of S. montelus was negatively correlated with latitude. These findings provided solid evidence to support the converse Bergmann's rule. We considered that the observed variation in morphological characteristics was most likely mediated by the seasonal length and thermoperiod to adapt to different latitudinal environment (e.g. shortened developmental time of immature stages for smaller body size at higher latitude). Moreover, the tendency towards progressively darker colour patterns (only in adult males) at increasingly low latitudes was consistent with Gloger's rule. We suggested that the observed colour variation was most likely associated with thermoregulation. Slight variation in the morphology of the W‐shaped stripe on the forewing of adult females was also found, and we presumed that the functions of sexual preferences, mimicry and thermoregulation might be involved.     

Revealing the colourful side of birds: spatial distribution of conspicuous plumage colours on the body of Australian birds

In many species of birds, different body parts often display very different colours. This spatial distribution of coloured plumage patches may be determined, among other factors, by the balance between being cryptic to predators, and conspicuous to intended receivers. If this is the case, ventral and anterior body parts in birds – which are less visible to predators but more prominent to conspecifics – should present more conspicuous and sexually dichromatic plumage colours. Here, I test these predictions using reflectance spectrometric measurements of standardised plumage patches across males and females for nearly an entire avifauna (Australian landbirds, n = 538 species). My data show that, as predicted, conspicuous and sexually dichromatic colours are mainly located near the head, while the plumage of the back is the most cryptic. One clear exception to this pattern is the conspicuous rump coloration. In many species, this patch can be concealed by wings, and therefore exposed only when necessary. In addition, conspicuous rump coloration could deflect or confuse predators in case of attack. However, there is considerable variation across species, and this makes position on the body a very poor predictor of plumage elaboration (R² < 0.02). Future studies should try to determine whether differences between species in the distribution of colours across the plumage are due to variation in ecological factors (predation risk, habitat, etc.).     

Spatial Variation in Fish Assemblages across small Mediterranean Drainages: Effects of Habitat and Landscape Context

Much debate about assemblage organization in stream fish may stem from analysing the effects of both local and large-scale processes on assemblage attributes over whole geographic regions. This study addresses this issue, by examining the contribution of local habitat attributes and landscape context to fish assemblage variation across small Mediterranean drainages in southern Portugal. Fish abundance and species composition was estimated in 28 sites, across 10 drainages, in both a dry year (1999) and a wet year (2001), and related to two sets of variables reflecting habitat and landscape characteristics. Fish showed responses to both sets of variables with variance partitioning indicating that landscape context had important effects on species richness whereas habitat attributes were the primary determinants of local fish abundance. In general, high species richness was associated with larger drainage area and higher rainfall variability, whereas variation in species abundances mostly reflected the influence of width, depth, conductivity, current velocity, substrate size and emergent vegetation. The relative contributions of both landscape context and habitat attributes to species richness and abundance were generally lower in 1999 than in 2001, with much less diversified species–habitat relationships being found in the former dry year. These results point to the dynamic nature of assemblage organization, emphasizing the importance of innovative, multi-scale approaches in advancing our knowledge of fish assemblage structure in Mediterranean streams.     

Half-Century Winter Duck Abundance and Temperature Trends in the Mississippi and Atlantic Flyways

Increasing concerns exist about possible decreased wintering duck abundance and hunting opportunities in the southern regions of the Atlantic and Mississippi flyways of North America. Researchers suggest these decreased abundances of ducks may be related to winter warming and related climatic phenomena. Accordingly, we tested predictions that duck abundance was increasing more at northern than southern latitudes, and that trends were related to average winter temperatures (Dec–Jan). We tested predictions using National Audubon Society Christmas Bird Count (CBC) data collected during December 1969 through January 2019 from 31 states in the United States and 6 Canadian provinces that comprise the Atlantic and Mississippi flyways for 16 species of dabbling and diving ducks (Anatinae). We found support for the prediction that CBC trends in duck abundance vary with latitude, and mean winter temperature explained nearly half the variation in CBC trends for 12 of 16 species. For some species, trends were negative in warmer regions and positive in colder regions. For others, trends were stable or slightly positive in warmer regions but more positive in colder regions. These results provide empirical evidence supporting climate-influenced winter range changes by important game duck species and suggest challenges and opportunities for waterfowl population, habitat, and hunting management in North America and the northern hemisphere. © 2021 The Wildlife Society.     

Quantitative genetics of a carotenoid-based color: heritability and persistent natal environmental effects in the great tit

The information content of signals such as animal coloration depends on the extent to which variation reflects underlying biological processes. Although animal coloration has received considerable attention, little work has addressed the quantitative genetics of color variation in natural populations. We investigated the quantitative genetics of a carotenoid-based color patch, the ventral plumage of mature great tits (Parus major), in a wild population. Carotenoid-based colors are often suggested to reflect environmental variation in carotenoid availability, but numerous mechanisms could also lead to genetic variation in coloration. Analyses of individuals of known origin showed that, although plumage chromaticity (i.e., color) was moderately heritable, there was no significant heritability to achromaticity (i.e., brightness). We detected multiple long-lasting effects of natal environment, with hatching date and brood size both negatively related to plumage chromaticity at maturity. Our reflectance measures contrasted in their spatiotemporal sensitivity, with plumage chromaticity exhibiting significant spatial variation and achromatic variation exhibiting marked annual variation. Hence, color variation in this species reflects both genetic and environmental influences on different scales. Our analyses demonstrate the context dependence of components of color variation and suggest that color patches may convey multiple aspects of individual state.