Melanism and diel activity of alpine Colias (Lepidoptera: Pieridae)

Summary The hypothesis that increased melanism provides a benefit in prolonging diel activity through more efficient absorption of solar radiation was tested in the field on a population of alpine Colias sulphur butterflies. A marked increase in the duration of flight and feeding behaviour existed for melanistic individuals when compared to lighter individuals under cool temperatures and low intensity solar radiation. More melanistic butterflies moved longer distance per day, and emigrated from the population at a faster rate. At high temperature and high radiant load lighter coloured individuals appeared more active. This is the first field demonstration of the advantage of melanism for increasing activity of ectotherms in cold environments.     

Influence of climate on the presence of colour polymorphism in two montane reptile species

The coloration of ectotherms plays an important role in thermoregulation processes. Dark individuals should heat up faster and be able to reach a higher body temperature than light individuals and should therefore have benefits in cool areas. In central Europe, montane local populations of adder (Vipera berus) and asp viper (Vipera aspis) exhibit a varying proportion of melanistic individuals. We tested whether the presence of melanistic V. aspis and V. berus could be explained by climatic conditions. We measured the climatic niche position and breadth of monomorphic (including strictly patterned individuals) and polymorphic local populations, calculated their niche overlap and tested for niche equivalency and similarity. In accordance with expectations, niche overlap between polymorphic local populations of both species is high, and even higher than that of polymorphic versus monomorphic montane local populations of V. aspis, suggesting a predominant role of melanism in determining the niche of ectothermic vertebrates. However, unexpectedly, the niche of polymorphic local populations of both species is narrower than that of monomorphic ones, indicating that colour polymorphism does not always enable the exploitation of a greater variability of resources, at least at the intraspecific level. Overall, our results suggest that melanism might be present only when the thermoregulatory benefit is higher than the cost of predation.     

Thermal melanism in ectotherms

Whether melanism plays a significant role in thermoregulation has been a persistent question in studies of thermal biology of ectotherms. This review provides a synthesis of the thermal melanism hypothesis which states that dark individuals (i.e. lower skin reflectance) are at an advantage under conditions of low temperature since they heat up faster than light individuals at a given level of solar radiation. Although skin color is an important trait in the thermal biology of ectotherms, it has rarely been explored in non-insect models. We draw on the available literature to assess the validity of four key assumptions that underlie this hypothesis. Ample support was found for the assumption that melanistic diurnal species inhabit cooler areas than lighter species and that melanism results in greater fitness in cold climates. By contrast, little direct support could be found for the assumption that there is a consistent melanism–body size tradeoff. Finally, the assumption that color, thermal physiology and behavior are coadapted has some support but requires further investigation. Overall, the functional, molecular and adaptive mechanisms of thermal melanism await further study.     

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.     

Variation in Melanism and Female Preference in Proximate but Ecologically Distinct Environments

Variation in pigmentation traits is an effective window to evolutionary processes due to their importance for survival and reproduction. In particular, one of the leading hypotheses for the maintenance of conspicuous pigmentation in natural populations is its signaling function in mate choice. Here, we demonstrate the occurrence of melanism in poeciliid fishes of the genus Poecilia that inhabits toxic, hydrogen sulfide springs in southern Mexico and the absence of melanism from closely related populations in reference habitats lacking hydrogen sulfide. Assays of female mate preference in both habitat types were used to examine whether divergence in female preference for melanism contributes to its maintenance in hydrogen sulfide springs. We found significant variation in female preferences for melanistic males. Specifically, melanistic females from the toxic spring exhibited a significant preference for melanistic males, while non‐melanistic females from the same population exhibited no preference. Females from the non‐sulfidic reference population discriminated strongly against melanistic males. Preferences of melanistic females appear to be a significant force in the maintenance of melanism in sulfidic habitats and suggest a change in preference as the divergence from non‐sulfidic ancestors. Potential polymorphism in preference within the hydrogen sulfide spring indicates that preference for melanistic males may not be environmentally controlled. Thus, a change in preference following divergence can promote the maintenance of variation in pigmentation within populations and between divergent habitats.     

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.     

Microstructure,evolution, and ontogeny of scale surfaces in cordylid and gerrhosaurid lizards

The distal oberhautchen and proximal beta layer of 41 species and every genus of cordylid and gerrhosaurid lizard were examined with a scanning electron microscope. Scale organs in these families are concentrated on the face, and scale organs occur singly or in pairs below the keel on the dorsals of cordylids but not gerrhosaurids. Most species examined possess primitive, lenticular scale organs. Unique and previously unknown, projecting scale organs occur in two species of Cordylus. Scale organs appear to be absent in species of Chamaesaura. The scale surfaces of cordylids are covered in macrohoneycomb. Individual compartments of cordylid macrohoneycomb are not intracellular structures but may be overlain by apparent oberhautchen cells. In some cordylid species, part of each cell is free and projects away from the scale surface as a pointed flap. Gerrhosaurids lack macrohoneycomb. Instead, their scale surfaces are covered in lamellae without dentate borders, 50–100 times as long as wide. The epidermis of Platysaurus is unique among cordylids in possessing lamellae 1–8 times as wide as long that overlap macrohoneycomb in some regions of the body. The scale surfaces of Cordylus cordylus undergo an ontogenetic change similar to that recently described for the xenosaurid genus Shinisaurus. Phylogenetic analysis of scale surface features allows the partitioning of Cordylus into three species groups and Chamaesaura into two species groups. Synapomorphies of the species of Chamaesaura, the species of Platysaurus, and the genera of the gerrhosaurids are also described. © 1995 Wiley-Liss, Inc.     

Molecular genetics and evolution of melanism in the cat family

Melanistic coat coloration occurs as a common polymorphism in 11 of 37 felid species and reaches high population frequency in some cases but never achieves complete fixation. To investigate the genetic basis, adaptive significance, and evolutionary history of melanistic variants in the Felidae, we mapped, cloned, and sequenced the cat homologs of two putative candidate genes for melanism (ASIP [agouti] and MC1R) and identified three independent deletions associated with dark coloration in three different felid species. Association and transmission analyses revealed that a 2 bp deletion in the ASIP gene specifies black coloration in domestic cats, and two different "in-frame" deletions in the MC1R gene are implicated in melanism in jaguars and jaguarundis. Melanistic individuals from five other felid species did not carry any of these mutations, implying that there are at least four independent genetic origins for melanism in the cat family. The inferred multiple origins and independent historical elevation in population frequency of felid melanistic mutations suggest the occurrence of adaptive evolution of this visible phenotype in a group of related free-ranging species.     

Epidermal glands in cordylid lizards,with special reference to generation glands

We investigated inter‐ and intraspecific variation in epidermal gland characteristics in cordylid lizards. Our particular interest was whether a causal relationship exists between the presence of generation glands and lifestyle, i.e. in rock‐dwelling or ground‐dwelling lizards. We established that both femoral and generation glands are always present in adult males, but that species can be divided into three categories on the basis of the presence of these glands in adult females: both femoral and generation glands absent; femoral glands present, but generation glands absent; and both femoral and generation glands present. The absence of epidermal glands in females appears to be the basal condition in the Cordylidae. All cordylid species are sexually dimorphic as far as generation gland number is concerned, with females having consistently less glands than males. Gland number in females is strongly affected by climate, being lower in cooler compared with warmer environments. In males, there is no clear geographical pattern, although in some clades males of high‐altitude species have more generation glands than males of species occurring at lower altitudes. At least four of the five ground‐dwelling species in the family display unique generation gland characteristics, compared with rock‐dwelling species, reflecting the continued importance of generation glands in terrestrial environments. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 312–324.     

Variation in the evolutionary integration of melanism with behavioral and physiological traits in <Emphasis Type="Italic">Xiphophorus variatus</Emphasis>

Vertebrate pigmentation is emerging as a powerful system for studying the evolution of adaptive traits and the maintenance of genetic and phenotypic variation in natural populations. Though melanism has been linked to physiological and behavioral traits in a variety of taxa, the generality of these associations for many taxa such as fishes remains unclear. Here I tested whether variation in melanism in a livebearing fish was correlated with a variety of traits often tested in other taxa: locomotor stress coping style during confinement, boldness in a novel environment, and metabolic rate. There were significant negative associations between an individual’s amount of melanistic pigmentation and both activity in confinement and boldness in a novel environment. In contrast with evidence from many prior studies, there was no relationship between melanism and metabolic rate. Overall, the data provide some support for documented relationships between melanism and behavioral traits, but did not support the generally reported relationship between melanism and metabolic rate. Links between melanism and behavioral coping strategies related to environmental stressors may have important implications for the evolution and maintenance of behavioral and morphological variation in natural populations. Nonetheless, these results also suggest variation among taxa in the extent to which pleiotropy has evolved between melanism and diverse organismal traits.     

不同经度地区北草蜥的喜好体温和热耐受性

在外温动物热生理特征的进化理论中,“静态”和“易变”是两个持续争论的对立观点。热生理学特征的种内变异是检验此类假设的最有力证据。本研究比较了不同经度地区北草蜥的热环境和热生理特征,以检验“静态”和“易变”假设。东部沿海地区(宁德)的环境温度高于内陆地区(贵阳),与之相适应,沿海地区北草蜥的喜好体温也高于内陆地区。然而,两地区蜥蜴的上临界温度和下临界温度无显著差异。尽管这些热生理学特征的种群间变异趋势并不一致,但是喜好温度随环境温度变化而改变的结果符合“易变”假设的预测。此外,本研究表明蜥蜴的喜好体温存在沿经度方向的地理变异。     

Thermal environment shapes cuticle melanism and melanin-based immunity in the ground cricket Allonemobius socius

The thermal melanin hypothesis posits that ectothermic individuals of larger size or from colder environments exhibit darker cuticles due to melanin’s efficacy in absorbing solar radiation. However, melanin is also a crucial component of arthropod immunity. Thus, thermal selection for increased cuticle darkness may profoundly influence melanin-based immune function. In this study, we address the relationships between the thermal environment (season length), cuticular melanism and two aspects of melanin-based immunity across nine thermally distinct populations of the cricket Allonemobius socius. We found that season length (i.e. degree days) and body size had a positive association with cuticle melanism in both sexes across populations, supporting the thermal melanism hypothesis. Despite their smaller size, males were found to have darker cuticles and superior melanin-based immunity. This pattern may be the result of additional selection on males due to sex-specific temperature-dependent activities, such as male calling song. Perhaps most interestingly, we found that short season length populations (i.e. colder) exhibited a greater phenoloxidase activity (aspect of the melanin-based immune system) in addition to darker cuticles in both sexes. This pattern is consistent with direct thermal selection on cuticular color, coupled with indirect selection on melanin-based immunity due to pleiotropy. Thus, thermal selection on cuticle darkness appears to indirectly shape the evolution of pathogen resistance in this system, and potentially for other terrestrial arthropod systems whose ranges encompass a significant thermal gradient.     

Plasticity and local adaptation explain lizard cold tolerance

How does climate variation limit the range of species and what does it take for species to colonize new regions? In this issue of Molecular Ecology, Campbell‐Staton et al. ( 2018 ) address these broad questions by investigating cold tolerance adaptation in the green anole lizard (Anolis carolinensis) across a latitudinal transect. By integrating physiological data, gene expression data and acclimation experiments, the authors disentangle the mechanisms underlying cold adaptation. They first establish that cold tolerance adaptation in Anolis lizards follows the predictions of the oxygen‐ and capacity‐limited thermal tolerance hypothesis, which states that organisms are limited by temperature thresholds at which oxygen supply cannot meet demand. They then explore the drivers of cold tolerance at a finer scale, finding evidence that northern populations are adapted to cooler thermal regimes and that both phenotypic plasticity and heritable genetic variation contribute to cold tolerance. The integration of physiological and gene expression data further highlights the varied mechanisms that drive cold tolerance adaptation in Anolis lizards, including both supply‐side and demand‐side adaptations that improve oxygen economy. Altogether, their work provides new insight into the physiological and genetic mechanisms underlying adaptation to new climatic niches and demonstrates that cold tolerance in northern lizard populations is achieved through the synergy of physiological plasticity and local genetic adaptation for thermal performance.     

Variation in thermally induced melanism in monarch butterflies (Lepidoptera: Nymphalidae) from three North American populations

• 1.As ectotherms, insects often experience varying temperatures throughout their life cycle, and some respond by becoming more or less melanistic (dark coloring) during development to increase or decrease thermal energy absorption as larvae or adults.
• 2.Monarch butterflies (Danaus plexippus) breed in temperate and tropical environments worldwide and are exposed to different average and extreme temperatures in different parts of their geographic range. In this study, we compared variation in thermally induced melanism among monarch butterflies from eastern and western North America and from South Florida.
• 3.We raised the progeny of wild-captured adult butterflies from these populations in a common garden experiment, rearing individuals in cold (19 °C), moderate (26 °C), and hot (32 °C) temperatures to examine population variation in larval and adult pigmentation.
• 4.Across all populations, monarch larvae developed the darkest coloration in the cold treatment and were lightest when reared in hot temperatures. Similar results were observed for measures of adult wing melanism, with the exception of adult females, which developed darker colored wings in warmer temperatures.
• 5.Significant population-level differences in average measures of melanism among larvae and adult butterflies were observed. Larvae from the eastern population became substantially darker in colder temperatures than S. Florida or western larvae. Western larvae were lightest overall, which might be adaptive to high temperatures experienced throughout portions of their summer breeding range. S. Florida larvae showed a lower response to cold temperatures relative to monarchs from either migratory population.
• 6.Population level differences were also observed for thermal responses in wing melanism, particularly among adult females. Moreover, we found significant family level effects for each measure of larval and adult melanism, pointing to a genetic basis or strong maternal effects influencing these traits in monarch butterflies.

     

EVOLUTION OF VIVIPARITY: A PHYLOGENETIC TEST OF THE COLD‐CLIMATE HYPOTHESIS IN PHRYNOSOMATID LIZARDS

The evolution of viviparity is a key life‐history transition in vertebrates, but the selective forces favoring its evolution are not fully understood. With >100 origins of viviparity, squamate reptiles (lizards and snakes) are ideal for addressing this issue. Some evidence from field and laboratory studies supports the “cold‐climate” hypothesis, wherein viviparity provides an advantage in cold environments by allowing mothers to maintain higher temperatures for developing embryos. Surprisingly, the cold‐climate hypothesis has not been tested using both climatic data and phylogenetic comparative methods. Here, we investigate the evolution of viviparity in the lizard family Phrynosomatidae using GIS‐based environmental data, an extensive phylogeny (117 species), and recently developed comparative methods. We find significant relationships between viviparity and lower temperatures during the warmest (egg‐laying) season, strongly supporting the cold‐climate hypothesis. Remarkably, we also find that viviparity tends to evolve more frequently at tropical latitudes, despite its association with cooler climates. Our results help explain this and two related patterns that seemingly contradict the cold‐climate hypothesis: the presence of viviparous species restricted to low‐elevation tropical regions and the paucity of viviparous species at high latitudes. Finally, we examine whether viviparous taxa may be at higher risk of extinction from anthropogenic climate change.     

Two new trypanosome species from East African cordylid lizards

Two species ofTrypanosoma are described from East African cordylid lizards.Trypanosoma zonuri n. sp. was found in a disjunct population ofCordylus cordylus on the western slope of Ngorongoro Crater in Arusha Region, northern Tanzania. It has the appearance of a delicate and fragile trypanosome, slightly over half the size of the robust, leaf-likeT. cordyli n. sp. that parasitisesCordylus t. tropidosternum in the Rondo Forest, Lindi Region, southern Tanzania. These are the first trypanosome species recorded from the saurian family Cordylidae in East Africa.     

Coloration affects heating and cooling in three color morphs of the Australian bluetongue lizard,Tiliqua scincoides

The color-mediated thermoregulation hypothesis predicts that dark body color (low reflectance) allows organisms to gain heat more efficiently than does pale coloration (high reflectance). This prediction is intuitive and widely assumed to be true, but has poor empirical support. We used rare, captive-bred, mutant melanistic, albino and wild-type Australian bluetongue lizards, Tiliqua scincoides to measure the effects of skin reflectance on the heating and cooling rates. We measured heating under an artificial radiant heat source and cooling rates in an ice-cooled box using live lizards in a room with still air. The effect of skin reflectance on heat transfer was clear, despite the substantial influence of body size. Melanistic T. scincoides showed low reflectance and gained heat faster than highly reflective albinos. Melanistic lizards also lost heat faster than albinos. Wild-type lizards were intermediate in reflectance, gained heat at rates indistinguishable from melanistic lizards, and lost heat at rates indistinguishable from albino lizards. This study system allowed us to control for variables that were confounded in other studies and may explain the inconsistent support for the color-mediated thermoregulation hypothesis. Our results provide clear evidence that skin reflectance influences the rate of heating and cooling in ectotherms.     

Macroecological Patterns of Climatic Niche Breadth Variation in Lacertid Lizards

Measuring climatic niche position and breadth may help to determine where species can occur over space and time. Using GIS-based and phylogenetic comparative methods, we investigated global patterns of variation in climatic niche breadth in lacertid lizards to test the following three hypotheses about climatic niche widths. First, does a species' temperature or precipitation niche breadth relate to its temperature or precipitation niche position(the mean value of annual mean temperature or annual precipitation across sampled localities in the range of each species)? Second, are there trade-offs between a species' temperature niche breadth and precipitation niche breadth? Third, does a species' temperature or precipitation niche breadth relate to altitude or latitude? We expect that:(1) species distributed in cold regions are specialized for low-temperature environments(i.e. narrow niche breadth center around low temperatures);(2) a negative relationship between species niche breadth on temperature and precipitation axes according to the tradeoff hypothesis(i.e. species that tolerate a broad range of precipitation regimes cannot also tolerate a broad range of temperatures);(3) precipitation niche breadth decreases with altitude or latitude, whereas temperature climatic niche breadth increases with altitude or latitude. Based on the analytical results we found that:(1) temperature niche breadth and position are negatively related, while precipitation niche breadth and position are positively related;(2) there is no trade-off between temperature and precipitation niche breadths; and (3) temperature niche breadth and latitude/altitude are positively related, but precipitation niche breadth and latitude/altitude are not significantly related. Our results show many similarities with previous studies on climatic niche widths reported for amphibians and lizards, which provide further evidence that such macroecological patterns of variation in climatic niche breadths may be widespread.     

Convergent evolution of embryonic growth and development in the eastern fence lizard (Sceloporus undulatus)

Theory predicts that cold environments will select for strategies that enhance the growth of ectotherms, such as early emergence from nests and more efficient use of resources. We used a common garden experiment to detect parallel clines in rates of embryonic growth and development by eastern fence lizards (Sceloporus undulatus). Using realistic thermal conditions, we measured growth efficiencies and incubation periods of lizards from five populations representing two distinct clades. In both clades, embryos from cold environments (Indiana, New Jersey, and Virginia) grew more efficiently and hatched earlier than embryos from warm environments (Florida and South Carolina). Because eggs from cold environments were larger than eggs from warm environments, we experimentally miniaturized eggs from one population (Virginia) to determine whether rapid growth and development were caused by a greater maternal investment. Embryos in miniaturized eggs grew as efficiently and incubated for the same duration as embryos in unmanipulated eggs. Taken together, our results suggest countergradient variation has evolved at least twice in S. undulatus.     

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.