INTEGRATING PHYLOGENY AND EXPERIMENTAL ETHOLOGY: FROM PATTERN TO PROCEs

Phylogenetic systematics offers ethologists a method for uncovering macroevolutionary patterns of behavioral diversity, which can be used to generate predictions for experimental investigation of microevolutionary processes. I used phylogenetic methods to disentangle the contributions of various selection pressures in the evolution of sexually dimorphic nuptial coloration in gasterosteid fishes in general, and Gasterosteus aculeatus in particular. Based upon the phylogenetic relationships between breeding colors and breeding behaviors in the gasterosteids, three predictions relevant to a discussion of nuptial coloration in G. aculeatus were proposed. The origin and elaboration of nuptial coloration was (1) weakly associated with male/male interactions (intrasexual selection), (2) moderately associated with parental care (natural selection) and (3) strongly associated with male/female interactions (intersexual selection). I tested the hypothesized macroevolutionary relationships between male breeding color and behavior in two sets of experiments studying changes in male nuptial coloration across the breeding cycle and female choice based on intensity of male color. The results of these experiments at the microevolutionary level of analysis corroborated the macroevolutionary predictions and confirmed the original intuition of previous researchers: male body color intensity is an important component of female mate choice in G. aculeatus. This study demonstrates that a phylogenetic hypothesis can generate predictions that are experimentally testable and falsifiable.     

Aggressive Behavior and Courtship Rejection in Brightly and Plainly Colored Female Keeled Earless Lizards (Holbrookia propinqua)

Relationships between bright secondary sexual coloration and behavior were studied in female Holbrookia propinqua, which develop striking orange and yellow colors during the breeding season. In tethered introduction studies, brightly colored females performed aggressive courtship rejection behaviors toward conspecific males; plainly colored females were not aggressive toward males, but attempted to avoid them. Responses of females of the two color patterns toward conspecific females of both color phases were not detectably different. Experimental introductions of lizards with coloration modified by paint showed that females of both color patterns recognize any other lizard bearing the bright female colors as female, regardless of actual sex. Both the orange and yellow components were shown to contribute to sex recognition. The yellow component alone allowed accurate sex identification, but only half the females responded to males painted with only the orange female component as if they were females. Because females did not behave differently toward other females on the basis of coloration, the hypothesis that bright coloration evolved as an adaptive signal between females is rejected. The dark ventrolateral stripes of male and plainly colored females did not appear to affect intraspecific social responses by females.     

Seasonal changes in blue tit crown color: do they signal individual quality?

Plumage coloration is generally perceived as a static traitand therefore not a good indicator of current condition. However,changing of feather colors after molt does occur and may haveimportant implications for signal function and sexual selection.We studied longitudinal changes in blue tit (Parus caeruleus)crown ultraviolet (UV)/blue color, a sexually selected trait,by repeatedly measuring the same individuals between early winterand late spring. Whereas crown UV reflectance (UV chroma andhue) decreased dramatically over time, brightness and saturationdid not show consistent patterns of change. The magnitude ofthe decline in coloration exceeded sexual and age dichromatismin hue and UV chroma, respectively. Hence, seasonal color changescould have strong effects on blue tit sexual signaling. Between-individualvariation in the decline in UV coloration was large and relatedto attributes of male, but not female, quality, such as sizeand condition. Thus, conspecifics could potentially gain informationabout male phenotypic quality by assessing color change overthe year. However, the degree of decline in male UV color didnot affect breeding success because neither the number of within-pairnor the number of extrapair offspring produced correlated withchanges in crown color. Seasonal changes in the expression ofplumage coloration are probably widespread, and maintainingplumage coloration could thus constitute an additional honesty-enforcingmechanism after molt is completed.     

Seasonal changes in the dorsal coloration in the lizard Aspidoscelis costata costata (Squamata: Teiidae)

Color and color patterns in animals are important characteristics that bring protection, by dampening the ability of predators that depend on their sight to detect their preys. In lizards, the dorsal coloration plays a key role in communication of intraspecific signals such as social cues. In this study, we evaluated the seasonal changes in the dorsal coloration of the wide foraging lizard A. costata costata, in Tonatico, State of Mexico, Mexico. The seasonal evaluation included: the rainy season from mid June to mid September (can also include the end of May to early October); and the dry season for the rest of the year. The dorsal coloration of A. costata costata and their microhabitats were evaluated by contrasting the color pattern with an identification guide and the control colors of Pantone, during 11 samplings carried out from February-October 2007. Individual lizard analysis recorded snout-vent length, sex and stage (juveniles and adults). Besides, all animals were marked by toe-clipping, allowing to distinguish dorsal coloration between seasons, sex and stage. A total of 95 lizards were analyzed (53 and 42 for the dry and rainy seasons respectively). We found that the dorsal coloration in A. costata costata varies seasonally and with microhabitats: during the dry season individuals show a brown coloration whereas during the rainy season becomes greener, as the background dominant vegetation color. The results of the present study suggest that: 1) the variation in dorsal coloration in A. costata costata plays an important role in the survival (by cryptic camouflage) of this widely foraging species; 2) the changes in the dorsal coloration of A. costata costata are individually expressed traits, since the coloration of the same lizard is either brown or green depending on the season; and 3) the cryptic functions of the dorsal coloration in widely foraging species have been largely underestimated. We discuss the possible influence of the changes in coloration in an habitat that changes drastically between both dry and rainy seasons.     

Eye camouflage and false eyespots: chaetodontid responses to predators

Synopsis The roles of eye camouflage and eyespots are examined within the genusChaetodon as are the various theories explaining the evolutionary significance of the brilliant colors. While eye camouflage is not common among reef fishes, 91% of the 90 species ofChaetodon, have eyemasks (82) or black heads (4). Eye camouflage occurs concomitantly with diurnal false eyespots in 45.5% (41 of 90) of the species. Diurnal false eyespots serve to misdirect attacks by predators and/or to advertise unpalatability. False eyespots are located on areas of the body which allow escape and survival following an attack. Data suggesting that predators learn about the undesirability of butterflyfishes are presented. Butterflyfishes are inactive at night, forage during the day and spawn at dusk. It is unlikely that nocturnal color changes are useful in conspecific interactions and are therefore believed to provide visual cues to potential predators. Nocturnal eyespots probably function to intimidate potential predators but could also remind them of unpalatability. The aggression release hypothesis (Lorenz 1962, 1966) to explain the brilliant coloration of chaetodontids is not supported because butterflyfish coloration changes and few species are territorial. The species recognition hypothesis (Zumpe 1965) is not supported by results of field experiments. The disruptive coloration hypothesis (Longley 1917) is rejected as a general explanation for poster coloration but does explain the prevalence of eyebars ofChaetodon spp. The aposematic hypothesis (Gosline 1965) is supported by morphology, behavior, a lack of predation and field observations. The possibility of Mullerian mimicry is suggested. It is concluded that the primary selective force behind chaetodontid coloration, particularly eyespots, has been predation and color patterns have evolved to minimize this threat.     

The Control of Color in Birds

SYNOPSIS. The colors of birds result from deposition of pigments—mainlymelanins and carotenoids—in integumentary structures,chiefly the feathers. The plumages of birds indicate their age,sex, and mode of living, and play important roles in camouflage,mating, and establishment of territories. Since feathers aredead structures, change of color of feathers is effected throughdivestment (molt) and replacement. The color and pattern ofa feather are determined by the interplay of genetic and hormonalinfluences prevailing in its base during regeneration. Mostbirds replace their feathers at least once annually. Some wearthe same kind of basic plumage all the time butothers alternatea basic and breeding plumage, either in one (the male) or bothsexes. Still others may have more than two molts, adding supplementalplumage at certain times in the plumage cycle. The varietiesof patterns of molt, the kinds of plumage, and the colors andpatterns of feathers among birds apparently are the result ofseveral kinds of selection pressures working through evolution.     

Phylogenetic Correlation Between Male Nuptial Color and Behavioral Responses to Color Across a Diverse and Colorful Genus of Freshwater Fish (Etheostoma spp., Teleostei: Percidae)

Sexual selection theory predicts that preferences in both sexes select for the elaboration of male nuptial coloration, with empirical evidence supporting these predictions. Empirical studies are often limited in their taxonomic inclusiveness, however, and typically do not examine how male and female preferences contribute to macroevolutionary patterns of male color variation across multiple lineages in a clade. This study examined color preferences in a group of dichromatic freshwater fishes known as darters (genus Etheostoma) that vary in the presence of male coloration. The strengths of attraction to black, blue, gray, and red models were tested in females of 18 species, in addition to males of five species. We found a positive association between the presence of red or orange on the body and the amount of time associating with red models, suggesting color variation is at least in part due to variation in female preferences between species. Males also spent more time associating with colors that most closely resembled conspecifics, suggesting that preferential responses to color in males also can contribute to the diversity of nuptial coloration in darters.     

Ultrastructural variation tune wing coloration of a moth Asota caricae Fabricius, 1775

Butterflies and moths develop highly ordered coloration in their wing for signal transmission. We have investigated the ultrastructural arrangement of wing coloration of a moth Asota caricae, applying light, optical polarized, and scanning electron microscopy, and spectrophotometry. The forewing of the moth is brown in color with a white spot at the center. The hindwing is golden yellow in color with many black patches in it. The ventral part of the forewing and dorsal hindwing share the similar color pattern. The ventral part of the hindwing has dull coloration in comparison to the dorsal one although the pattern remains same. The spectrometry analysis reveals various patterns of absorbance and reflectance spectra for various colors. The peak observed for various colors remain same although the intensity of peak changes. Bright colors possess highly ordered structures whereas irregular structures are found in dull colored scales. The color variation observed due to dorsal and ventral part of the wing is due to the minute difference observed in terms of ultrastructural arrangement revealed by scanning electron microscope. The color pattern of A. caricae is due to variation of microstructures present within the scale.     

马毛色遗传机理研究进展

动物毛色是人类正向选择产生的表型之一,在遗传与进化过程中扮演着重要角色。其中马的毛色丰富多变,单从表型无法准确判别其属于哪种毛色,造成马品种登记时毛色性状记录不准确,因此研究马毛色形成机理在育种工作中具有重要意义。随着基因组学及测序技术的日益成熟,马毛色形成遗传机理的研究不断深入,并发现不同毛色性状与特定疾病之间的相关性。本文从遗传学的角度对马的毛色进行归类,对与其形成的相关基因、作用机理及应用等研究进展进行了综述,以期为马毛色形成机理的系统性研究和马匹选育提供借鉴和参考。     

Evolution of plumage coloration in the crow family (Corvidae) with a focus on the color-producing microstructures in the feathers: a comparison of eight species

Plumage coloration has been the subject for a variety of questions that comprise the center of modern evolutionary biology. Unlike carotenoids that the concentration directly influences the intensity of the color, melanin, in addition to produce brown or black colors, is often involved in producing the structural coloration such as glossiness or iridescence. As the melanin granules can be located in the barbs or the barbules, we aim to (i) discern if the colors observed at macro scale comes from the barbs, the barbules or both in a series of related species and (ii) estimate the evolutionary history of the color-producing mechanisms in the family Corvidae that are known to have melanin-based coloration. From a preliminary comparative analysis on eight representative species, we found three coloration schemes in Corvidae; (1) matte colors of brown or black that were produced in barbs and barbules; (2) non-iridescent structural colors such as blue, bluish gray and white, that were produced in the barbs and (3) iridescent structural colors that were produced only in distal barbules. Comparative character analysis of these coloration schemes suggests that the ancestral state among these species were the colors produced in the barbs and that the color produced in the distal barbules is a derived character. The evolution of iridescence seems tightly linked to the evolution of the colors produced in the distal barbules. Data from more species should be incorporated in order to grasp a full picture on the evolutionary history of plumage coloration in this group of birds.     

Cryptic plumage signaling in Aphelocoma Scrub-Jays

Recent studies of avian vision and plumage coloration have revealed a surprising degree of cryptic sexual dimorphism, with many examples of male–female differences in UV reflectance that are invisible to humans. We examined the potential for male–female and adult–subadult differences in plumage coloration in the genus Aphelocoma. This group of jays comprises 10 phylogenetic species, which are found across southern and central North America and include cooperatively breeding species, as well as species that form socially monogamous pair-bonds typical of most species of birds. Our goal was to determine whether male–female and adult–subadult differences in plumage coloration were more common in species with complex social systems (i.e., cooperative breeders). We collected a series of reflectance measurements from hundreds of museum specimens and analyzed the results using a model of an avian visual system. We found that age- and sex-related differences were not more frequent in species that practice cooperative breeding. Hence, plumage signaling relating sex and age may not be strongly associated with complex social systems. Rather, the relative lack of a stable and familiar social environment, as well as other selective pressures and constraints (e.g., habitat use and plumage complexity), may have favored a greater degree of age- and sex-related differences in plumage coloration in jays that practice simple biparental care.     

基于花青素代谢培育蓝色花卉的研究进展

花色是植物吸引昆虫传播花粉的主要因素,对于植物在自然界的生存必不可少,也是观赏植物最重要的性状之一。在蓬勃发展的花卉产业中,色彩各异花卉的培育,可以弥补自然花色的匮乏,但是令人垂涎的蓝色花比较难培育。花色的多样性主要是由花青素及其衍生物的种类和含量等因素决定的,飞燕草色素的合成是形成蓝色花的关键因素,许多植物体内缺少合成飞燕草色素的结构基因。近年来,利用基因工程技术培育蓝色花的研究也时有报道。文中以常见的观赏植物为例,基于花青素代谢调控,从影响飞燕草色素合成的关键因素和不同分子改良途径培育蓝色花等几个方面对植物花朵呈色的机制进行了综述,并展示不同分子育种策略可能在其他领域的应用,为其他植物或经济作物的色泽改良如彩色棉蓝色纤维的培育等提供参考和技术支持。     

Natural selection on social signals: signal efficacy and the evolution of chameleon display coloration

Whether general patterns of signal evolution can be explained by selection for signal efficacy (detectability) has yet to be established. To establish the importance of signal efficacy requires evidence that both signals and their detectability to receivers have evolved in response to habitat shifts in a predictable fashion. Here, we test whether habitat structure has predictable effects on the evolution of male and female display coloration in 21 lineages of African dwarf chameleon (Bradypodion), based on a phylogenetic comparative analysis. We used quantitative measures of display coloration and estimated signal detectability as the contrast of those colors among body regions or against the background vegetation as perceived by the chameleon visual system. Both male and female display colors varied predictably with different aspects of habitat structure. In several (but not all) instances, habitat-associated shifts in display coloration resulted in habitat-associated variation in detectability. While males exhibit a remarkable variety of colors and patterns, female display coloration is highly conserved, consisting in all populations of contrasting dark and light elements. This color pattern may maximize detectability across all habitat types, potentially explaining female conservatism. Overall, our results support the view that selection for signal efficacy plays an important role in the evolution of animal signals.     

Ornamental non-carotenoid red feathers of wild burrowing parrots

Bird plumage colors have the potential to indicate individual quality, condition, health, immunocompetence, or the extend of parental care. Color intensity of feathers has been found to correlate with parameters of individual quality, condition, parental care and breeding success. Psittaciformes are well known for their colorful plumage but the significance of parrot coloration is still poorly understood. Red colors are very common in many parrot species. They are produced by at least four non-carotenoid-based pigments (linear polyenal structure). In the present study, we investigated a collection of red abdominal feathers of a marked population of wild Burrowing Parrots Cyanoliseus patagonus in Patagonia, Argentina. The aims of this study were to investigate the ecological significance of the recently described non-carotenoid-based red pigments of Psittaciformes, and the relationships between objectively assessed plumage color and body size, body condition, breeding success and nestling growth in wild Psittaciformes. We found that sexes differed in plumage coloration (sexual dichromatism), that plumage color was a good predictor of female body condition and male size, and we identified the red coloration of the abdominal patch as a signal of individual quality and parental investment.     

Spectral Sensitivities and Color Signals in a Polymorphic Damselfly

Animal communication relies on conspicuous signals and compatible signal perception abilities. Good signal perception abilities are particularly important for polymorphic animals where mate choice can be a challenge. Behavioral studies suggest that polymorphic damselflies use their varying body colorations and/or color patterns as communication signal for mate choice and to control mating frequencies. However, solid evidence for this hypothesis combining physiological with spectral and behavioral data is scarce. We investigated this question in the Australian common blue tail damselfly, Ischnura heterosticta, which has pronounced female-limited polymorphism: andromorphs have a male-like blue coloration and gynomorphs display green/grey colors. We measured body color reflectance and investigated the visual capacities of each morph, showing that I. heterosticta have at least three types of photoreceptors sensitive to UV, blue, and green wavelength, and that this visual perception ability enables them to detect the spectral properties of the color signals emitted from the various color morphs in both males and females. We further demonstrate that different color morphs can be discriminated against each other and the vegetation based on color contrast. Finally, these findings were supported by field observations of natural mating pairs showing that mating partners are indeed chosen based on their body coloration. Our study provides the first comprehensive evidence for the function of body coloration on mate choice in polymorphic damselflies.     

White Ibis integument color during the breeding season

ABSTRACT.   Many birds undergo bare part color changes during the breeding season. Most investigators have focused on color as a signal of individual quality. An alternative, but not exclusive, function of bare part color may be signaling readiness to breed, especially in colonial, asynchronous breeders. White Ibises ( Eudocimus albus ) are colonial waterbirds that show vivid bare part colors on their bills and legs during reproduction. We quantified bill and leg colors to describe color changes and their possible relationship to reproductive status during the breeding season of White Ibises in the Florida Everglades from 1998 to 2001. We also examined the correlation between bare part colors and circulating concentrations of sex steroids to understand the factors that regulate bare part colors. During the display stage, male and female ibises developed dark pink bills and scarlet legs. As the breeding season progressed, bills and legs faded and developed a muted pink hue. The bare part colors of female ibises were correlated with testosterone concentrations, but those of male ibises were not correlated with any hormones. A discriminant function analysis based on principal component scores (representing variation in saturation and hue) and the amount of black on the bill successfully classified ibis reproductive stage 94% of the time. The use of bare part colors to determine reproductive status may be useful for studying reproduction in colonially nesting birds, where access to breeding sites can be difficult and potential for researcher disturbance is high.     

Melanin coloration in New World orioles II: ancestral state reconstruction reveals lability in the use of carotenoids and phaeomelanins

Although many animals use carotenoids to produce bright yellow, orange, and red colors, an increasing number of studies have found that other pigments, such as melanins, may also be used to produce bright colors. Yet, almost nothing is known about the evolutionary history of this colorful melanin use. We used reflectance spectrometry to determine whether colors in New World orioles were predominantly due to carotenoids, colorful melanins, or a mixture of both. We then used ancestral state reconstruction to infer the directionality of any pigment changes and to test for phylogenetic signal. We found that three oriole taxa likely switched from carotenoid- to melanin-based colors. Several other oriole taxa apparently gained localized melanin coloration, or had coloration that seemed to be produced by a mixture of carotenoids and melanins. We also found little phylogenetic signal on the use of carotenoids or melanins to produce color. However, all pigment changes occurred within one of three major clades of the oriole genus, suggesting there may be signal at deeper phylogenetic levels. These repeated independent switches between carotenoid and melanin colors are surprising in light of the important signaling role that color pigments (especially carotenoids) are thought to play across a wide range of taxa.     

UV plumage color is an honest signal of quality in male budgerigars

Elaborate and colorful feathers are important traits in female mate choice in birds. Plumage coloration can result from pigments deposited in feathers such as carotenoids and melanins, or can be caused by nano-scale reflective tissues (structurally based coloration), usually producing ultraviolet (UV) coloration. Structural colorations remain the least studied of the three most important feather colorations. Previous studies have found a female preference for UV color in the budgerigar, Melopsittacus undulatus, but it is not clear what information this ornament conveys, nor what is the possible cost associated with its production. We investigated possible correlations between immune response and plumage color of wild-type (green) male budgerigars. In particular we measured the wing web swelling resulting from injection of phytohaemagglutinin (PHA). We did not detect any correlation between the sedimentation rate and morphological and color variables. We found that UV chroma is the best predictor for the cutaneous immune activity. Indeed, male budgerigars with high UV reflectance in the breast feathers showed stronger immune responses. These results are consistent with the idea that UV colors are special signals conveying information about a bird’s condition.     

A test of an antipredatory function of conspicuous plastron coloration in hatchling turtles

Bright colorations in animals are sometimes an antipredatory signal meant to startle, warn, or deter a predator from consuming a prey organism. Freshwater turtle hatchlings of many species have bright ventral coloration with high internal contrast that may have an antipredator function. We used visual modeling and field experiments to test whether the plastron coloration of Chrysemys picta hatchlings deters predators. We found that bird predators can easily distinguish hatchling turtles from their backgrounds and can easily see color contrast within the plastron. Raccoons cannot easily discriminate within-plastron color contrast but can see hatchlings against common backgrounds. Despite this, we found that brightly-colored, high contrast, replica turtles were not attacked less than low contrast replica turtles, suggesting that the bright coloration is not likely to serve an antipredatory function in this context. We discuss the apparent lack of innate avoidance of orange coloration in freshwater turtles by predators and suggest that preference and avoidance of colors are context-dependent. Since the bright colors are likely not a signal, we hypothesize that the colors may be caused by pigments deposited in tissue from maternal reserves during development. In most species, these pigments fade ontogenetically but they may have important physiological functions in species that maintain the bright coloration throughout adulthood.     

Aposematic coloration does not deter corallivory by fish on the coral <Emphasis Type="Italic">Montastraea cavernosa</Emphasis>

Predation on corals by visual predators is a significant source of partial or total mortality on coral reefs, and corals have evolved strategies, including chemical defenses, to deter predation. One mechanism that organisms use to communicate the presence of chemical defenses is aposematic coloration, or the display of bright coloration as a warning to visual predators such as fish. Corals exhibit multiple colors, and it has been hypothesized that one role for this variability in coloration is as an aposematic warning of adverse palatability. Here, we test green and orange color morphs of the Caribbean coral Montastraea cavernosa for the presence of chemical defenses and whether their differences in coloration elicited different feeding responses. While M. cavernosa is chemically defended, there is no difference in feeding deterrence between color morphs; thus, the different color morphs of this coral species do not appear to represent an example of aposematic coloration.