Polymorphism and phene geography of the blue rock pigeon in Europe

The variation of blue rock pigeon plumage color has been studied in 192 settlements of Europe. As in earlier studies, six color phenotypes have been distinguished, the main of which are blue, intermediate and melanistic. The phenotype frequency distribution patterns in urban and rural landscapes have been determined. Pigeon populations with increased density are the most melanistic. Rural populations are less melanistic than urban ones. The frequency of birds with aberrant plumage colors varies randomly and is increased in some localities. The phenotypic structure of synanthropic populations of the blue rock pigeon in Europe displays a latitudinal gradient.     

Dynamics of balanced polymorphism morphs in blue rock pigeon (Columba livia) from Moscow

Analysis of the 30-year dynamics (from 1978 to 2008) of phenotype frequencies in the Moscow blue rock pigeon population revealed changes in the frequency distribution of several plumage color phenotypes. The frequency of the melanistic phenotype decreased as a result of the decrease in the total population size and, consequently, population density in colonies. This decrease in the pressure of the melanistic phenotype led to increased proportions of wild-type and aberrantly colored birds throughout the town and an increased frequency of transitory phenotypes in the central region. A predominance of one phenotype suppressed genetic diversity of plumage color phenotypes. The changes in the phenotype distribution in populations were associated with changes in the social structure of the human population. As the feeding resources grew poorer because of social anthropogenic factors, the pigeon population size decreased. This decrease in population size changed the phenotype frequencies, which was considered to be an adaptive response of the population to environmental changes.     

Dynamics of balanced polymorphism morphs in blue rock pigeon <Emphasis Type="Italic">Columbia livia</Emphasis>

Analysis of the 30-year dynamics (from 1978 to 2008) of phenotype frequencies in the Moscow blue rock pigeon population revealed changes in the frequency distribution of several plumage color phenotypes. The frequency of the melanistic phenotype decreased as a result of the decrease in the total population size and, consequently, population density in colonies. This decrease in the pressure of the melanistic phenotype led to increased proportions of wild-type and aberrantly colored birds throughout the town and an increased frequency of transitory phenotypes in the central region. A predominance of one phenotype suppressed genetic diversity of plumage color phenotypes. The changes in the phenotype distribution in populations were associated with changes in the social structure of the human population. As the feeding resources grew poorer because of social anthropogenic factors, the pigeon population size decreased. This decrease in population size changed the phenotype frequencies, which was considered to be an adaptive response of the population to environmental changes.     

Geographic variation in color of the synanthropic Blue rock pigeon

Variation in plumage color was studied in 211 populations of the blue rock pigeon (Columbia livia) from various geographic zones. The frequency distribution of three common and three rare phenotypes and an interspecific hybridization zone of C. livia and C. rupestris were described. Different conditions of the origin of the urban populations of blue rock pigeon and consequent high genetic heterogeneity of this species were revealed. The key aspect of color polymorphism in blue rock pigeon is synanthropization, i.e., anthropogenic evolution.     

Geographic Variation of Color in the Synanthropic Blue Rock Pigeon

Variation in plumage color was studied in 211 populations of the blue rock pigeon (Columbia livia) from various geographic zones. The frequency distribution of three common and three rare phenotypes and an interspecific hybridization zone ofC. liviaand C. rupestriswere described. Different conditions of the origin of the urban populations of blue rock pigeon and consequent high genetic heterogeneity of this species were revealed. The key aspect of color polymorphism in blue rock pigeon is synanthropization, i.e., anthropogenic evolution.     

Rapid evolution of fire melanism in replicated populations of pygmy grasshoppers

Evolutionary theory predicts an interactive process whereby spatiotemporal environmental heterogeneity will maintain genetic variation, while genetic and phenotypic diversity will buffer populations against stress and allow for fast adaptive evolution in rapidly changing environments. Here, we study color polymorphism patterns in pygmy grasshoppers (Tetrix subulata) and show that the frequency of the melanistic (black) color variant was higher in areas that had been ravaged by fires the previous year than in nonburned habitats, that, in burned areas, the frequency of melanistic grasshoppers dropped from ca. 50% one year after a fire to 30% after four years, and that the variation in frequencies of melanistic individuals among and within populations was genetically based on and represented evolutionary modifications. Dark coloration may confer a selective benefit mediated by enhanced camouflage in recently fire-ravaged areas characterized by blackened visual backgrounds before vegetation has recovered. These findings provide rare evidence for unusually large, extremely rapid adaptive contemporary evolution in replicated natural populations in response to divergent and fluctuating selection associated with spatiotemporal environmental changes.     

Pigeons in the sun: Thermal constraints of eumelanic plumage in the rock pigeon (Columba livia)

In wild vertebrates, several species exhibit eumelanic color polymorphism with the coexistence of dark and light morphs. The maintenance of such polymorphism suggests the existence of a selective balance between the morphs and a large body of literature has reported the costs and benefits of darker plumage coloration in birds. Among them, it has been suggested that melanin and dark plumage could entail high energetic costs especially under hot and sunny climates. However, to my knowledge, the thermal constraints of sun exposure have rarely been studied in polymorphic species. Here, we tested the impact of eumelanic plumage coloration on plumage and body temperatures, and evaporative cooling behavior in the polymorphic rock pigeon (Columbia livia). We experimentally exposed light and dark pigeons to direct sun radiation for 1 h while a few birds were maintained in the shade as controls. We found that sun exposure was associated with increased plumage temperature, and this effect was greater for darker pigeons. In addition, we found that sun exposure was also associated with higher cloacal temperature but for dark pigeons only. Finally, light and dark pigeons were more likely to show cooling evaporative behavior when exposed to sun and as their cloacal temperature increases. Altogether, these results suggest that darker pigeons may have a lower ability to cope with heat and solar radiations and that dark plumage can be associated with thermal costs in this polymorphic species.     

Reduced predation risk for melanistic pygmy grasshoppers in post‐fire environments

The existence of melanistic (black) color forms in many species represents interesting model systems that have played important roles for our understanding of selective processes, evolution of adaptations, and the maintenance of variation. A recent study reported on rapid evolutionary shifts in frequencies of the melanistic forms in replicated populations of Tetrix subulata pygmy grasshoppers; the incidence of the melanistic form was higher in recently burned areas with backgrounds blackened by fire than in nonburned areas, and it declined over time in postfire environments. Here, we tested the hypothesis that the frequency shifts of the black color variant were driven, at least in part, by changes in the selective regime imposed by visual predators. To study detectability of the melanistic form, we presented human “predators” with images of black grasshoppers and samples of the natural habitat on computer screens. We demonstrate that the protective value of black coloration differs between burnt and nonburnt environments and gradually increases in habitats that have been more blackened by fire. These findings support the notion that a black color pattern provides improved protection from visually oriented predators against blackened backgrounds and implicate camouflage and predation as important drivers of fire melanism in pygmy grasshoppers.     

Dispatches from the neighborhood watch: Using citizen science and field survey data to document color morph frequency in space and time

Heritable color polymorphisms have a long history of study in evolutionary biology, though they are less frequently examined today than in the past. These systems, where multiple discrete, visually identifiable color phenotypes co‐occur in the same population, are valuable for tracking evolutionary change and ascertaining the relative importance of different evolutionary mechanisms. Here, we use a combination of citizen science data and field surveys in the Great Lakes region of North America to identify patterns of color morph frequencies in the eastern gray squirrel (Sciurus carolinensis). Using over 68,000 individual squirrel records from both large and small spatial scales, we identify the following patterns: (a) the melanistic (black) phenotype is often localized but nonetheless widespread throughout the Great Lakes region, occurring in all states and provinces sampled. (b) In Ohio, where intensive surveys were performed, there is a weak but significantly positive association between color morph frequency and geographic proximity of populations. Nonetheless, even nearby populations often had radically different frequencies of the melanistic morph, which ranged from 0% to 96%. These patterns were mosaic rather than clinal. (c) In the Wooster, Ohio population, which had over eight years of continuous data on color morph frequency representing nearly 40,000 records, we found that the frequency of the melanistic morph increased gradually over time on some survey routes but decreased or did not change over time on others. These differences were statistically significant and occurred at very small spatial scales (on the order of hundreds of meters). Together, these patterns are suggestive of genetic drift as an important mechanism of evolutionary change in this system. We argue that studies of color polymorphism are still quite valuable in advancing our understanding of fundamental evolutionary processes, especially when coupled with the growing availability of data from citizen science efforts.     

Camouflage versus running performance as strategies against predation in a lizard inhabiting different habitats

Running speed and camouflage are associated with the foraging and anti‐predator abilities of animals. The toad‐headed lizard, Phrynocephalus versicolor, has evolved a darker dorsal color in melanistic habitats and maintained a lighter color in adjacent, non‐melanistic habitats. We test the hypothesis that lizards have weaker running speed on well‐matching backgrounds than on less matching backgrounds. We used lizard models to compare the predation pressure, while the running speed of dark and light lizards were compared in field tunnels using a video recording method. Our results indicated that both the dark lizards in melanistic Heishankou (HSK) and the light lizards in non‐melanistic Guazhou (GZ) face lower predation pressure than potential color‐background unmatched lizards. The light lizards have a potentially higher running speed than darker lizards in melanistic habitats, which implies that substrate color matching populations with benefits of camouflage might have lower anti‐predation pressure, and the costs of investment in melanin production may reduce running capacity.     

Sequence variation in the coding region of the melanocortin-1 receptor gene (MC1R) is not associated with plumage variation in the blue-crowned manakin (Lepidothrix coronata)

Avian plumage traits are the targets of both natural and sexual selection. Consequently, genetic changes resulting in plumage variation among closely related taxa might represent important evolutionary events. The molecular basis of such differences, however, is unknown in most cases. Sequence variation in the melanocortin-1 receptor gene (MC1R) is associated with melanistic phenotypes in many vertebrate taxa, including several avian species. The blue-crowned manakin (Lepidothrix coronata), a widespread, sexually dichromatic passerine, exhibits striking geographic variation in male plumage colour across its range in southern Central America and western Amazonia. Northern males are black with brilliant blue crowns whereas southern males are green with lighter blue crowns. We sequenced 810 bp of the MC1R coding region in 23 individuals spanning the range of male plumage variation. The only variable sites we detected among L. coronata sequences were four synonymous substitutions, none of which were strictly associated with either plumage type. Similarly, comparative analyses showed that L. coronata sequences were monomorphic at the three amino acid sites hypothesized to be functionally important in other birds. These results demonstrate that genes other than MC1R underlie melanic plumage polymorphism in blue-crowned manakins.     

The evolution of black plumage from blue in Australian fairy‐wrens (Maluridae): genetic and structural evidence

Genetic variation in the melanocortin‐1 receptor (MC1R) locus is responsible for color variation, particularly melanism, in many groups of vertebrates. Fairy‐wrens, Maluridae, are a family of Australian and New Guinean passerines with several instances of dramatic shifts in plumage coloration, both intra‐ and inter‐specifically. A number of these color changes are from bright blue to black plumage. In this study, we examined sequence variation at the MC1R locus in most genera and species of fairy‐wrens. Our primary focus was subspecies of the white‐winged fairy‐wren Malurus leucopterus in which two subspecies, each endemic to islands off the western Australian coast, are black while the mainland subspecies is blue. We found fourteen variable amino acid residues within M. leucopterus, but at only one position were alleles perfectly correlated with plumage color. Comparison with other fairy‐wren species showed that the blue mainland subspecies, not the black island subspecies, had a unique genotype. Examination of MC1R protein sequence variation across our sample of fairy‐wrens revealed no correlation between plumage color and sequence in this group. We thus conclude that amino acid changes in the MC1R locus are not directly responsible for the black plumage of the island subspecies of M. leucopterus. Our examination of the nanostructure of feathers from both black and blue subspecies of M. leucopterus and other black and blue fairy‐wren species clarifies the evolution of black plumage in this family. Our data indicate that the black white‐winged fairy‐wrens evolved from blue ancestors because vestiges of the nanostructure required for the production of blue coloration exist within their black feathers. Based on our phylogeographic analysis of M. leucopterus, in which the two black subspecies do not appear to be each other's closest relatives, we infer that there have been two independent evolutionary transitions from blue to black plumage. A third potential transition from blue to black appears to have occurred in a sister clade.     

Blue,not UV,plumage color is important in satin bowerbird Ptilonorhynchus violaceus display

Several studies have suggested that peak plumage reflectance in birds matches color preferences used in mate choice. We tested this hypothesis in adult satin bowerbird males that have a short‐wavelength saturated blue‐black plumage with a peak reflectance in the UV. We found that the chroma of the blue (405–480 nm), but not the peak reflecting UV (320–400 nm) portion of the male plumage spectrum was significantly correlated with male mating success. A plot of correlation coefficients between male mating success and plumage saturation showed a well‐defined peak in the blue. This suggests that: 1) blue plumage coloration is more important in mate choice than UV or other colors, and 2) that there is a mismatch between the peak reflectance of the plumage of male satin bowerbirds and the range of plumage wavelengths that are correlated with male mating success. This indicates that it is not safe to infer a role of UV or other colors in mate choice simply because of a peak in plumage reflectance.     

Albino and melanistic genets (<Emphasis Type="Italic">Genetta genetta</Emphasis>) in Europe

While melanistic genets (Genetta spp.) are well known in a few species of the genus, albino specimens have not been described to date. Here, we report on the presence of unusual albino and melanistic common genets (Genetta genetta) in Spain, discussing their frequency of occurrence in the wild. Melanistic and albino common genets are not known in the original African range of the species, thus phenotypical variability in coat colour appears to be greater in Europe, its introduced range. Natural (e.g. a reduced risk of predation in Europe) and/or artificial (e.g. captive-rearing of the species) selection could explain this fact, but more research on the topic is needed.     

Testing homology of loci for two plumage colors, "lavender" and "recessive white," with chicken and Japanese quail hybrids

Homology for two plumage color loci was studied by hybridization between chickens and Japanese quail. First, chicken-quail hybrids were produced from homozygous "lavender" chicken cocks and "bleu" Japanese quail, and all 30 hybrids had the same parental slate blue plumage color. On the other hand, no hybrids with this plumage were obtained out of 18 progeny from the same cocks and wild-type quail. These results show that the slate blue plumage color is determined by homologous loci in Japanese quail and chickens. Second, all (n = 25) chicken-quail hybrids hatched from homozygous "recessive white" cocks and "recessive white" (n = 8) or "wild-type" (n = 17) quail had the same pattern of plumage color, with white feathers on the ventral face and colored feathers elsewhere. These results indicate that the recessive white mutations are not homologous in Japanese quail and chickens.     

Opposing fitness consequences of colour pattern in male and female snakes

Local populations of the adder, Vipera berus, are polymorphic for dorsal colour pattern, containing both melanistic (black) and zig-zag patterned individuals. Colour patterns in snakes influence crypsis and thermoregulatory capacity and therefore may be subjected to natural selection. To find an explanation for the maintenance of this polymorphism I examined temporal and spatial variation in morph frequency, and tested for differential selection among morphs using data from a six year capture-mark-recapture study. The data derive from six groups of islands in the Baltic Sea off the Swedish east coast, two mainland localities near the coast, and one inland locality. Morph frequency did not change over time within a population but varied among populations: melanistic individuals were not found at the inland locality, but comprised from 17 to 62% of the coastal and island populations. Adders frequently moved between islands within a group, but the tendency to disperse was independent of morph. These results suggest that the polymorphism is stable and maintained by a deterministic process. Scar frequency was twice as high among melanistic as among zig-zag snakes, and melanistic individuals were easier to capture, indicating that predation may be higher on the melanistic morph. Colour morphs did not differ in body size, but analysis of recapture data shows evidence for differential survival among morphs. Zig-zag males survived better than melanistic males, but the relative survival rates of morphs were reversed in females. This difference was consistent through time and may be due to sexual differences in behaviour, with melanism increasing predation intensity when associated with male but not with female behaviour. Opposing fitness consequences of colour pattern in the two sexes may help maintain colour polymorphism within populations of Vipera berus.     

Plumage color as a status signal in male–male interaction in the red-flanked bushrobin, Tarsiger cyanurus

Recent studies have suggested that structural-based coloration is an honest signal of male genetic and/or conditional quality in sexual selection. However, whether structural coloration functions in intrasexual competition is unknown. We examined whether plumage color functions as a status signal during intrasexual interactions in the red-flanked bushrobin Tarsiger cyanurus; adult males have many blue plumes as structural coloration whereas yearling males and females are olive brown with few blue plumages. Blue males did not always dominate olive-brown males. The number of interactions did not differ with the colors of the two birds involved. The interactions of a blue male and an olive-brown male were less aggressive than those of two blue or of two olive-brown males. In this study, we found that structural plumage coloration may serve as a signal of aggressive intent and lower the escalation level of an aggressive interaction in a manner consistent with hypotheses regarding the evolution of delayed plumage maturation.     

Evolution of avian plumage color in a tetrahedral color space: a phylogenetic analysis of new world buntings

We use a tetrahedral color space to describe and analyze male plumage color variation and evolution in a clade of New World buntings--Cyanocompsa and Passerina (Aves: Cardinalidae). The Goldsmith color space models the relative stimulation of the four retinal cones, using the integrals of the product of plumage reflectance spectra and cone sensitivity functions. A color is represented as a vector defined by the relative stimulation of the four cone types--ultraviolet, blue, green, and red. Color vectors are plotted in a tetrahedral, or quaternary, plot with the achromatic point at the origin and the ultraviolet/violet channel along the Z-axis. Each color vector is specified by the spherical coordinates theta, phi, and r. Hue is given by the angles theta and phi. Chroma is given by the magnitude of r, the distance from the achromatic origin. Color vectors of all distinct patches in a plumage characterize the plumage color phenotype. We describe the variation in color space occupancy of male bunting plumages, using various measures of color contrast, hue contrast and diversity, and chroma. Comparative phylogenetic analyses using linear parsimony (in MacClade) and generalized least squares (GLS) models (in CONTINUOUS) with a molecular phylogeny of the group document that plumage color evolution in the clade has been very dynamic. The single best-fit GLS evolutionary model of plumage color variation over the entire clade is a directional change model with no phylogenetic correlation among species. However, phylogenetic innovations in feather color production mechanisms--derived pheomelanin and carotenoid expression in two lineages--created new opportunities to colonize novel areas of color space and fostered the explosive differentiation in plumage color. Comparison of the tetrahedral color space of Goldsmith with that of Endler and Mielke demonstrates that both provide essentially identical results. Evolution of avian ultraviolet/violet opsin sensitivity in relation to chromatic experience is discussed.     

Why cordylid lizards are black at the south-western tip of Africa

There is a high incidence of melanism among the cordylid lizards of Africa and it was suggested that melanism has a thermoregulatory function in these species. One prediction of the thermal melanism hypothesis would be that melanistic cordylid species would be restricted to cool environments. The aim of our study was to determine the climatic conditions with which melanistic cordylids are currently associated. Distributional data for the eight melanistic cordylid taxa occurring at the south-western tip of Africa were obtained from the CapeNature State of Biodiversity database. Using GIS and principal component analyses, the climatic variables best describing the geographical distribution of melanistic cordylids were investigated. We found that several melanistic cordylid populations show a distinct association with a high incidence of fog and cloud cover, underscoring the thermal melanism hypothesis. Several other populations, however, do not show such an association. We conclude that some species, such as Pseudocordylus capensis , have a morphology that provides them with great flexibility in habitat use, allowing them to overcome the constraints of melanism in warm environments. The available information suggests that melanism in cordylids evolved during the Miocene in response to the development of the cold Benguela Current along the west coast of Africa. We also conclude that viviparity, a sit-and-wait foraging strategy and a rock-dwelling lifestyle would have preadapted cordylids for survival in cool conditions brought about by the development of the cold Benguela Current. These conditions would have been unfavourable for oviparous, active-foraging lizards. The co-occurrence of melanistic lizards and melanistic molluscs at the Landdroskop locality casts some doubt on improved heating rates as the only explanation for melanism in cordylids, because molluscs are not heliothermic baskers.     

The molecular basis of an avian plumage polymorphism in the wild: a melanocortin-1-receptor point mutation is perfectly associated with the melanic plumage morph of the bananaquit, Coereba flaveola

BACKGROUND: Evolution depends on natural selection acting on phenotypic variation, but the genes responsible for phenotypic variation in natural populations of vertebrates are rarely known. The molecular genetic basis for plumage color variation has not been described in any wild bird. Bananaquits (Coereba flaveola) are small passerine birds that occur as two main plumage variants, a widespread yellow morph with dark back and yellow breast and a virtually all black melanic morph. A candidate gene for this color difference is the melanocortin-1 receptor (MC1R), a key regulator of melanin synthesis in feather melanocytes. RESULTS: We sequenced the MC1R gene from four Caribbean populations of the bananaquit; two populations of the yellow morph and two populations containing both the yellow morph and the melanic morph. A point mutation resulting in the replacement of glutamate with lysine was present in at least one allele of the MC1R gene in all melanic birds and was absent in all yellow morph birds. This substitution probably causes the color variation, as the same substitution is responsible for melanism in domestic chickens and mice. The evolutionary relationships among the MC1R haplotypes show that the melanic alleles on Grenada and St. Vincent had a single origin. The low prevalence of nonsynonymous substitutions among yellow haplotypes suggests that they have been under stabilizing selection, whereas strong selective constraint on melanic haplotypes is absent. CONCLUSIONS: We conclude that a mutation in the MC1R is responsible for the plumage polymorphism in a wild bird population and that the melanic MC1R alleles in Grenada and St. Vincent bananaquit populations have a single evolutionary origin from a yellow allele.