Habitat light, colour variation, and ultraviolet reflectance in the Grand Cayman anole, Anolis conspersus

Abstract: Colour patches are complex traits, the components of which may evolve independently through a variety of mechanisms. Although usually treated as simple, two-dimensional characters and classified as either structural or pigmentary, in reality colour patches are complicated, three-dimensional structures that often contain multiple pigment types and structural features. The basic dermal chromatophore unit of fishes, reptiles and amphibians consists of three contiguous cell layers. Xanthophores and erythrophores in the outermost layer contain carotenoid and pteridine pigments that absorb short-wave light; iridophores in the middle layer contain crystalline platelets that reflect light back through the xanthophores; and melanophores in the basal layer contain melanins that absorb light across the spectrum. Changes in any one component of a chromatophore unit can drastically alter the reflectance spectrum produced, and for any given adaptive outcome (e.g. an increase in visibility), there may be multiple biochemical or cellular routes that evolution could take, allowing for divergent responses by different populations or species to similar selection regimes. All of the mechanisms of signal evolution that previously have been applied to single ornaments (including whole colour patches) could potentially be applied to the individual components of colour patches. To reach a complete understanding of colour patch evolution, however, it may be necessary to take an explicitly multi-trait approach. Here, we review multiple trait evolution theory and the basic mechanisms of colour production in fishes, reptiles and amphibians, and use a combination of computer simulations and empirical examples to show how multiple trait evolution theory can be applied to the components of single colour patches. This integrative perspective on animal colouration opens up a host of new questions and hypotheses. We offer specific, testable functional hypotheses for the most common pigmentary (carotenoid, pteridine and melanin) and structural components of vertebrate colour patches.

Abstract: We tested the prediction of the sensory drive hypothesis using four allopatric populations of the lizard Anolis cristatellus from two distinct environments (i.e., mesic and xeric conditions). For each population, we measured habitat light characteristics and quantified signal design by measuring the spectral and total reflectance and transmittance of the dewlap. We used these data to calculate dewlap detectability using an empirically based model of signal detection probability. We found that populations from mesic and xeric conditions occupy two distinct habitats with respect to light intensity and spectral quality and that dewlap design has diverged between populations in a way that increases signal detectability in each habitat. The major difference in dewlap design was in total reflectance and transmittance, making dewlaps from xeric habitats darker and dewlaps from mesic habitats brighter. Furthermore, dewlap detection decreased significantly when a dewlap from a xeric habitat is detected u...

Abstract: Effective visual communication requires signals that are easy to detect, transmit, receive, and discriminate. Animals can increase the probability that their visual signals would be detected by evolving signals that contrast with their visual background. Animals can further enhance this contrast by behaviorally modifying the existing visual background. Male golden-collared manakins (Manacus vitellinus) clear leaf litter from the ground to form courts, which are used as display arenas. Using reflectance measures of the signal (male plumage) and the visual background (cleared court and adjacent litter), the irradiance measures of ambient light during display, and published measures of photoreceptor sensitivity of a Passerine, we test the hypothesis that court-clearing augments the contrast between male plumage and the visual background. We find that the chromatic and brightness contrasts of golden patches used during courtship are greater against the cleared court than against adjacent litter. In addition, we find that cleared courts provide a less variable background for these color patches, resulting in displays that consistently contrast the visual background. These results suggest that behavioral modification of the visual background may act to increase the conspicuousness of colorful male plumage during display, providing an explanation for why golden-collared manakins, and possibly other species, build or clear display courts. Key words: chromatic contrasts, court-clearing, Manacus vitellinus, manakins, signaling, visual signals. [Behav Ecol 15:1003–1010 (2004)]

Abstract: Many animal species display striking color differences with respect to geographic location, sex, and body region. Traditional adaptive explanations for such complex patterns invoke an interaction between selection for conspicuous signals and natural selection for crypsis. Although there is now a substantial body of evidence supporting the role of sexual selection for signaling functions, quantitative studies of crypsis remain comparatively rare. Here, we combine objective measures of coloration with information on predator visual sensitivities to study the role of crypsis in the evolution of color variation in an Australian lizard species complex (Ctenophorus decresii). We apply a model that allows us to quantify crypsis in terms of the visual contrast of the lizards against their natural backgrounds, as perceived by potential avian predators. We then use these quantitative estimates of crypsis to answer the following questions. Are there significant differences in crypsis conspicuousness among populations? Are there significant differences in crypsis conspicuousness between the sexes? Are body regions "exposed" to visual predators more cryptic than "hidden" body regions? Is there evidence for local adaptation with respect to crypsis against different substrates? In general, our results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations. Body regions that varied most in contrast between the sexes and between populations were also most conspicuous and are emphasized by males during social and sexual signaling. However, results varied with respect to the aspect of coloration studied. Results based on chromatic contrast ("hue" of color) provided better support for the crypsis hypothesis than did results based on achromatic contrast ("brightness" of color). Taken together, these results support the view that crypsis plays a substantial role in the evolution of color variation and that color patterns represent a balance between the need for conspicuousness for signaling and the need for crypsis to avoid predation.

Abstract: Potential prey are often exposed to multiple predators that vary in their foraging tactics and ability to detect prey. For animals that rely on crypsis to avoid predators, one solution is to alter their behaviour or appearance to maximize crypsis in ways that are specific to different types of predator. We tested whether dwarf chameleons (Bradypodion transvaalense) showed different behavioural responses, including colour change, towards multiple predators (bird and snake models) that detect and capture prey in different ways, and whether these antipredator responses varied geographically. Chameleons consistently used the same body postures (lateral compression and flipping to the opposite side of the branch) and displayed similar chromatic (colour) contrast against the natural background in response to both predator types. However, they became significantly more achromatically contrasting (brighter) in the presence of the snake compared to the bird. This relative difference in achromatic contrast towards the two types of predator was consistent among populations. There were also significant differences in both absolute achromatic and chromatic contrast among populations despite very similar light environment, background coloration and habitat structure. Our results highlight facultative crypsis as one type of flexible antipredator tactic and emphasize the importance of visual ecology in understanding prey–predator interactions. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88, 437–446.

Abstract: There is a bewildering diversity of signals, sensory systems, and signaling behavior. A consideration of how these traits affect each other's evolution explains some of this diversity. Natural selection favors signals, receptors, and signaling behavior that maximize the received signals relative to background noise and minimize signal degradation. Properties of sensory systems bias the direction of evolution of the signals that they receive. For example, females may prefer males whose signals they can perceive more easily, and this will lead to the spread of more easily perceived male traits. Environmental conditions during signal transmission and detection also affect signal perception. Specific environmental conditions will bias the evolutionary direction of behavior, which affects the time and place of signaling as well as microhabitat preferences. Increased specialization of microhabitats and signaling behavior may lead to biased evolution of the sensory systems to work more efficiently. Thus, sensory...

Abstract: Sympatric native Anolis species with similar structural habitats but contrasting climatic habitats are closer in head and body size on species—rich than on depauperate islands. In two localities, sympatric Anolis species with differential occurrences in sun or shade sought lower, more shaded perches during midday, resulting in partly nonsynchronous utilization of the vegetation by the two species. The second observation may be related to the first in the following way: nonsynchronous spatial overlap could dictate relatively great resource overlap for species coinhabiting patchy or edge areas, requiring great differences between the species in prey size in addition to those in climatic habitat. The extent of such overlap on small depauperate islands could be greater if these contained a greater proportion of patchy or edge habitats (with respect to insolation), or if climatic preferences were broader and more overlapping than on large, species—rich islands. In each locality, the relatively more shade—inhab...

Abstract: All too often in evolutionary biology we are led to speculate or infer the mode of action of natural selection; we usually do not know why some individuals are more adaptive than others. Very often attempts to measure natural selection are unsuccessful, leading to heated arguments about' the relative importance of selection, genetic drift, and epistasis in evolution (Lewontin, 1974). Until we know more about how and why natural selection occurs, attempts to measure it are quixotic, and discussions of its importance are theandric. It is no coincidence that most of the successful studies of natural selection have dealt with animal color patterns; it should be obvious which color patterns are more adaptive in the presence of visually hunting predators. The adaptive significance of warning coloration and mimicry of distasteful species has been worked out (Cott, 1940; Wickler, 1968; Edmunds, 1974; Rothschild, 1975; Turner, 1977). But most species are neither distasteful nor mimetic; most have inconspicuous or cryptic color patterns in their natural habitats (Poulton, 1890; Thayer, 1909; Cott, 1940; Endler, 1978). Most field and experimental studies have shown that the overall color or tone of inconspicuous species matches or approximates the background (DiCesnola, 1904; Sumner, 1934, 1935; Isley, 1938; Popham, 1942; Dice, 1947; Kettlewell, 1956, 1973; Turner, 1961; Kaufman, 1974; Wicklund, 1975; Curio, 1976), but they treated species with solid colors or

Abstract: In studies of animal colouration it is no longer necessary to rely on subjective assessments of colour and conspicuousness, nor on methods which rely upon human vision. This is important because animals vary greatly in colour vision and colour is context-dependent. New methods make it practical to measure the colour spectrum of pattern elements (patches) of animals and their visual backgrounds for the conditions under which patch spectra reach the conspecific's, predator's or prey's eyes. These methods can be used in both terrestrial and aquatic habitats. A patch's colour is dependent not only upon its reflectance spectrum, but also upon the ambient light spectrum, the transmission properties of air or water, and the veiling light spectrum. These factors change with time of day, weather, season and microhabitat, so colours must be measured under the conditions prevalent when colour patterns are normally used. Methods of measuring, classifying and comparing colours are presented, as well as techniques for assessing the conspicuousness of colour patterns as a whole. Some implications of the effect of environmental light and vision are also discussed.

Abstract: It has long been known that the general colors and tones of animals tend to match their backgrounds (E. Darwin, 1794; Poulton, 1890). The adaptive significance of this has been borne out in numerous experimental studies (DiCesnola, 1904; Sumner, 1934, 1935; Isley, 1938; Popham, 1942; Dice, 1947; Turner, 1961; Kettlewell, 1956, 1973; Kaufman, 1974; Wiklund, 1975; Curio, 1976). There is also a good understanding of warning coloration (Cott, 1940; Wickler, 1968; Edmunds, 1974; Rothschild, 1975). However, the determinants of color pattern are poorly known, although it is known in a general way that the patterns and forms of animals are similar to their backgrounds (Poulton, 1890; Thayer, 1909; Cott, 1940; Wickler, 1968; Robinson, 1969; Edmunds, 1974; Fogden and Fogden, 1974). It is the purpose of this paper to explore the factors that determine color patterns under various specific conditions. The basic assumption is that a color pattern must resemble a random sample of the background seen by predators in order to be cryptic, and must deviate from the background in one or more ways in order to be conspicuous. As a result, the actual pattern evolved in a particular place represents a compromise between factors which favor crypsis and those which favor conspicuous color patterns.