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.     

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.     

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.     

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.     

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.     

Animal visual systems and the evolution of color patterns: sensory processing illuminates signal evolution

Abstract Animal color pattern phenotypes evolve rapidly. What influences their evolution? Because color patterns are used in communication, selection for signal efficacy, relative to the intended receiver's visual system, may explain and predict the direction of evolution. We investigated this in bowerbirds, whose color patterns consist of plumage, bower structure, and ornaments and whose visual displays are presented under predictable visual conditions. We used data on avian vision, environmental conditions, color pattern properties, and an estimate of the bowerbird phylogeny to test hypotheses about evolutionary effects of visual processing. Different components of the color pattern evolve differently. Plumage sexual dimorphism increased and then decreased, while overall (plumage plus bower) visual contrast increased. The use of bowers allows relative crypsis of the bird but increased efficacy of the signal as a whole. Ornaments do not elaborate existing plumage features but instead are innovations (new color schemes) that increase signal efficacy. Isolation between species could be facilitated by plumage but not ornaments, because we observed character displacement only in plumage. Bowerbird color pattern evolution is at least partially predictable from the function of the visual system and from knowledge of different functions of different components of the color patterns. This provides clues to how more constrained visual signaling systems may evolve.     

Red cell enzyme types in rheumatic diseases

We studied the frequencies of red cell enzyme types, AcP, PGM1 and EsD in 213 patients with rheumatoid arthritis (RA), systemic lupus erythematosus (SLE), rheumatic heart disease (RHD), scleroderma (Scl) and psoriatic arthropathy (PsA). The differences in frequency of AcP phenotypes between RA, Scl, and PsA and the Moscow population were significant. In PsA the PGM1 phenotype 1-1 frequency was significantly decreased, while the phenotype 2-1 frequency was significantly increased.     

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.     

Ecologically conditioned changes in gene frequencies for polymorphic systems in the nonnative population of the northeastern USSR]

Examination of 1051 healthy residents of Magadan in terms of nine polymorphic systems has shown changes in phenotype frequencies depending on duration of the persons' staying in the North. The frequency pattern of phenotypes is significantly different among men and women. The phenotype frequencies of alkaline serum phosphatase and acidic erythrocyte phosphatase were found in a more strict correlation with the northern residence term. The part played by selective migratory behaviour in development of genetic structure of population is discussed and also possible ecologic factors resulting in phenotype and gene frequency changes are analysed.     

Costs and Benefits of Subadult Plumage in Mute Swans: Testing Hypotheses for the Evolution of Delayed Plumage Maturation

In some avian species, young birds capable of reproducing diminish their prospects of doing so by molting into a subadult plumage that accurately signals their subadult status. Several hypotheses have been proposed to explain the evolution of delayed plumage maturation, but testing them usually has involved interspecific comparisons that are hard to interpret. Mute swans (Cygnus olor) exhibit two phenotypes that differ in whether the birds have a gray subadult plumage (SAP phenotype) or molt immediately into an all white adult plumage (AP phenotype). The AP phenotype results from a recessive allele on the X chromosome; both phenotypes occur in the same population and even in the same brood. We compared costs and benefits of both phenotypes in mute swans on the Chesapeake Bay in 1972-1980 and on Long Island Sound in 1982-1989. Swans with the SAP phenotype had higher survival rates from hatching to fledging than AP swans. In the fall, when AP cygnets began to molt into their white plumage, their parents often attacked and drove them off while allowing SAP cygnets from the same brood to remain on their territories for several more months. SAP males had higher survival rates during their first 2 yr of life than AP males, but AP swans bred at a younger age than SAP swans. The only proposed hypothesis for the evolution of delayed plumage maturation that can explain its occurrence in mute swans is the status-signaling hypothesis. This hypothesis argues that males with subadult plumage honestly advertise their age and subordinate status while AP swans are cheaters and engaging in dishonest communication. SAP males acquire a longer period of parental care, suffer less aggression from older birds, and increase their survival but forgo the opportunity to breed at an early age. This is a unique example of how a single gene resulted in either honest or dishonest communication, changed a bird's relationship with its parents and potential mates, and altered the bird's chances to survive and to reproduce.     

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.     

Are long‐term widespread avian body size changes related to food availability? A test using contemporaneous changes in carotenoid‐based color

Recent changes in global climate have been linked with changes in animal body size. While declines in body size are commonly explained as an adaptive thermoregulatory response to climate warming, many species do not decline in size, and alternative explanations for size change exist. One possibility is that temporal changes in animal body size are driven by changes in environmental productivity and food availability. This hypothesis is difficult to test due to the lack of suitable estimates that go back in time. Here, we use an alternative, indirect, approach and assess whether continent‐wide changes over the previous 100 years in body size in 15 species of Australian birds are associated with changes in their yellow carotenoid‐based plumage coloration. This type of coloration is strongly affected by food availability because birds cannot synthesize carotenoids and need to ingest them, and because color expression depends on general body condition. We found significant continent‐wide intraspecific temporal changes in body size (wing length) and yellow carotenoid‐based color (plumage reflectance) for half the species. Direction and magnitude of changes were highly variable among species. Meta‐analysis indicated that neither body size nor yellow plumage color showed a consistent temporal trend and that changes in color were not correlated with changes in size over the past 100 years. We conclude that our data provide no evidence that broad‐scale variation in food availability is a general explanation for continent‐wide changes in body size in this group of species. The interspecific variability in temporal changes in size as well as color suggests that it might be unlikely that a single factor drives these changes, and more detailed studies of museum specimens and long‐term field studies are required to disentangle the processes involved.     

First description of color variations in the annual killifish Millerichthys robustus,and preliminary observations about its geographical distribution

Millerichthys robustus is the only annual killifish distributed in America with phenotypic color variations, not yet described. Accordingly, we first describe the color pattern in both sexes to define its phenotypical variations. We then analyze the frequency of these phenotypes on a geographical scale, in four localities that represent opposite points of Millerichthys’s distribution in the Mexican southeast. Color analysis based on the RGB system allowed us to define five-color phenotypes in males continuously distributed in various perceptual units between two extreme colors (yellow-red): yellow, moderate orange, dark orange, strong orange and red. These color patterns found in M. robustus could be attributed to melanin, carotenoid, and pteridine pigments. The orange phenotypes was present in all localities studied. The yellow phenotype was present only in northeastern and northwestern locations, and the red phenotype was present only in northern populations. Female color variations were observed in the number of ocelli (from 1 to 15) at the base of the caudal peduncle and dorsal region. Ocelli have been associated with anti-predator functions because they resemble the eyes of vertebrates, thus shifting the target of predator attacks to less vital body parts. Females with 3 ocelli were the most frequent phenotype, and females with 13–15 ocelli occurred only in the northern populations. We concluded that male and female of M. robustus are not randomly distributed along their distribution range, which suggest that color phenotypes may react differently to biotic and abiotic factors that probably determine their distribution and frequency within the studied population.

     

Genetics of Ustilago violacea. XXXII. Genetic evidence for transposable elements

Crosses between Ustilago violacea mutant strains with different color phenotypes that were derived from the 1.A1 and 2.A2 laboratory strains yielded, as expected, bisectored teliospore colonies with the parental colors as well as the a-1 and the a-2 mating-types. Generally, wild teliospore collections usually produced sporidia of both mating-types, providing two-mating-type (TMT) strains. Occasionally, however, sporidia with only one mating-type allele, a-1 or a-2, were obtained from teliospores, providing one-mating-type (OMT) strains. Crosses between OMT and laboratory strains with different color phenotypes gave (1) bisectored teliospore colonies with the parental colors or colonies with a parental color and a nonparental color and (2) nonsectored colonies with the nonparental color or with the parental color. The frequencies for the occurrence of non-parental color ranged from 41% to 93%, depending on the strain. The yield of teliospore colonies was usually reduced for these crosses. In many of these teliospore colonies, morphologically-altered sporidia (MAS phenotype) were observed. The morphology and the size of the sporidia with the MAS phenotype differed from those of teliospore colonies of the crosses between the laboratory strains. In addition, these sporidia did not form conjugants. A cross involving the TMT strains C449 yielded the MAS phenotype as well as a high incidence of tetrad colonies with a nonparental color. The high degree of instability of the parental color phenotypes, and the high frequency of the appearance of nonparental color phenotypes as well as the appearance of the MAS phenotype, are in accord with the presence of active and inactive transposable elements in the OMT strains, TMT strains, and laboratory strains.     

Haptoglobin polymorphism among Saharian and West African groups. Haptoglobin phenotype determination by radioimmunoelectrophoresis on Hp O samples.

The haptoglobin (Hp) polymorphism is investigated in 11 African groups living in an area from the Algerian Sahara to Central Africa. More than 4,000 samples were examined. In the Saharian samples, the Hp1 gene frequency is higher than in any other African group. From north to south, a decrease in the Hp1 gene frequency is observed; in the Pygmy sample only, this frequency is lower than the frequency of the Hp2 gene. By means of a sensitive radioimmunoelectrophoresis, the presence of a residual Hp in Hp O sera in which the Hp polymorphism can also be determined can be revealed. Absence of Hp 1-1 and significant excess of Hp 2-2 individuals were observed. More Hp 2-1M phenotypes were detected in the Hp O population than in the non-Hp O population examined. In the Hp O samples, the influence of the phenotype distribution on the Hp gene frequencies is discussed. The heavy polymers of the Hp related to the presence of the alpha 2 chain (Hp2 gene product) are involved only in the biological mechanisms responsible for the presence of Hp O and Hp 2-1 M phenotypes among African groups.     

Evolutionarily stable sex ratios and sex allocations

The paternal fitness of a sexual individual is equated with the fitness of those eggs of its potential mates which it is able to fertilize. This property enables the total sexual fitness of individuals to be expressed in terms of female gamete contributions in separate equations for a cosex (an individual in a population composed of a single sexual class which combines male and female functions) and for parents in a dioecious population. The general equations are used in phenotypic models of selection which examine conditions maximizing the fitness advantage of one phenotype over another with a different sex ratio or allocation. As an example, it is shown that finite population size confers full stability on the sexual allocations in a cosexual population and on the sex ratio in a dioecious population.The use of fitness advantages provides the outcome of selection for all frequencies of contrasted phenotypes. It is therefore possible to redefine an ESS to allow for persistent variability in a population. A phenotype is an ESS in a population if, from any initial frequency, it is protected from loss by its fitness advantage. The conditions for a rare mutant to spread invariably coincide with those for its fixation only if an individual of any phenotype affects the fitness of other individuals of all phenotypes in identical ways.     

Individual responses to population size structure: the role of size variation in controlling expression of a trophic polyphenism

We investigated how size structure affects development of alternative larval phenotypes in Arizona tiger salamanders, Ambystoma tigrinum nebulosum, by testing the hypothesis that population size structure per se is a significant component of an individual's environment. Larvae of this subspecies exhibit one of two feeding phenotypes; typical larvae eat zooplankton and macroinvertebrates and cannibalistic larvae feed primarily on conspecifics. Previous laboratory experiments showed that larval density positively affected expression of the cannibalistic phenotype. In this study we tested the hypothesis that size variation among larvae also serves as a cue triggering development of the cannibalistic phenotype. We report laboratory experiments and field observations showing that both an individual larva's position in a size distribution and the amount of size vaiation among larvae serve as cues stimulating development of cannibalistic larvae. Larval density and population size structure provide a larva with an indication of the abundance and vulnerability of potential conspecific prey. Size variation among larvae, in turn, appears to be influenced by larval density. Thus, a complex relationship exists between larval density, population size structure, and the frequency of cannibals within a habitat.     

Production of plumage ornaments among males and females of two closely related tropical passerine bird species

The evolution of elaborate secondary sexual traits (i.e., ornaments) is well‐studied in males but less so in females. Similarity in the appearance of ornaments between males and females supports the view that female ornaments arise as a neutral byproduct of selection on male traits due to genetic correlation between sexes, but recent research suggests an adaptive function of female ornaments in at least some contexts. Information on the degree to which production of ornaments differs between the sexes can shed light on these alternative perspectives. We therefore characterized the structural underpinnings of melanin‐based plumage production in males and females of two closely related passerine bird species (genus Malurus). Importantly, both ornamented and unornamented phenotypes in each sex are present between these two species, providing an opportunity to test the null expectation of equivalent modes of production in male and female ornamented phenotypes. In Malurus alboscapulatus, ornamented females are qualitatively similar to males, but we describe a distinctive ornamented female phenotype that differs from that of males in lacking a blue sheen and in lower feather barbule density. In M. melanocephalus, unornamented males and females are also similar in appearance, and we describe a similarity between unornamented phenotypes of males and females in both color and underlying feather barbule structure and pigment composition. Unornamented male M. melanocephalus can flexibly transition to the ornamented phenotype in weeks, and we found extreme differences in color and feather structure between these two alternative male phenotypes. These results contradict the idea that female ornaments have evolved in this system following a simple switch to male‐like plumage by demonstrating greater complexity in the production of the ornamented phenotype in males than in females.