Dissertation (Ph.D.)--Auburn University, 2005. / Abstract. Includes bibliographic references.

Alpine melanism, a tendency to become darker with increased elevation and latitude, occurs in many species of butterflies. In Parnassius phoebus (Papilionidae) it has been assumed that alpine melanism is the result of adaptation to colder environments at higher elevations and latitudes, owing to the ability of darker wings to transfer more solar energy to the body as heat. I examined five independently variable components of wing melanism for agreement with this hypothesis, as well as the subsidiary hypothesis that size is inversely correlated to melanism because it also affects thermoregulation. Basal patch blackness (proportion of scales which are black) in both sexes is well correlated with elevation, latitude, and air temperature during the flight season. In males, increased basal patch blackness is advantageous because it increases the proportion of total time spent in flight looking for mates, but is locally disadvantageous because it increases the probability of dispersal. Basal patch width is only poorly correlated with elevation, latitude, and air temperature during the flight season in males, and not correlated in females. Increased patch width also results in increased flight activity, but does not affect dispersal. Basal wing transparency has little correlation with elevation, and no effect on activity, although it does slightly affect body temperature. Within individual mountains, distal wing blackness and transparency are correlated with elevation, but show no correlation when many mountains are considered together. Distal wing characters have no effect on flight activity or dispersal. Size is well correlated with elevation and latitude, but is less well correlated with air temperature. Smaller males disappear from the population more rapidly than large males, apparently through increased mortality because small males do not disperse more rapidly. Females disperse more than males, with larger females dispersing more than smaller females. Variation in basal patch blackness may have a similar adaptive significance in both sexes, but all the other melanism characters appear to have a different significance for each sex. I develop a new hypothesis for the mechanism of the effect of wing color on thermoregulation in dorsal basking butterflies. I suggest that most transfer of solar radiation to the body by the wings occurs while in flight, rather than while dorsal basking on a substrate. This hypothesis is used to explain the observation that the minimum levels of air temperature and solar radiation at which flight can occur does not depend on wing color in Parnassius phoebus, in contrast to phenotype dependent flight thresholds in lateral basking Colias. / Science, Faculty of / Zoology, Department of / Graduate

Explaining the wide range of animal colouration in the natural world is a key issue in evolutionary biology. Bumble bees are often brightly coloured and show a range of colours and colour patterns in different species as well as considerable variation within species. The large garden bumble bee, Bombus ruderatus, is highly variable in its degree of black (melanic) colouration, with morphs ranging from the familiar yellow and black bands (banded) through intermediate forms to morphs that are totally melanic. The aim of this research was to determine what might be maintaining the colour polymorphism in populations of B. ruderatus in the South Island, New Zealand. Colouration of worker bees was measured using a digital photography method and found to be significantly different across sample sites. To look at potential adaptive functions of body colour in B. ruderatus, three hypotheses of thermoregulation, desiccation tolerance and Müllerian mimicry were tested by comparing patterns of variation in melanism to patterns of variation in climatic variables (temperature, rainfall, humidity) and abundance of conspecifics. In order to address the possibility that selectively neutral processes were more important than selection, the genetic structure of B. ruderatus populations was characterised and compared to the pattern of variation in melanism. The colouration of individuals from the same population collected at different times in the season was compared to evaluate whether body colour was plastic and any support for the genetic basis of melanism in B. ruderatus was also assessed by determining any relationship between relatedness and degree of melanism. The results suggest that differences in the degree of melanism between populations are greater than the differences expected through selectively neutral forces alone and, therefore, that the pattern of variation in melanism is likely a result of selection and/or phenotypic plasticity in addition to gene flow and genetic drift. Although a global model consisting of four climatic variables and the abundance of conspecifics explained a small proportion of the variation in melanism, no support was found for any specific hypothesis relating to the adaptive function for body colour. Instead the results suggest that some combination of factors, most likely including factors not measured in this study, is influencing the frequency of melanic morphs. In addition, there was evidence that body colour was influenced by phenotypic plasticity and that melanism has a low heritability in B. ruderatus. Taken together, these results imply that patterns of melanism across B. ruderatus populations are complex and it is likely that multiple factors are influencing melanism in concert.

Many insect populations become darker at high elevations and high latitudes. Despite absence of empirical evidence, it is commonly believed that melanism allows more efficient basking by insects in sunlight, thereby raising body temperature and increasing activity. Variation in melanism within a single population of alpine Colias sulphur butterflies (Lepidoptera:Pieridae) allowed determination of the advantage, in cold environments, for this characteristic. Alternative hypotheses relating the effect of melanism to fecundity, predation, diel activity, and seasonal survivorship were tested by field and laboratory observation and experiment. At low temperature, melanistic females are more fecund. A balancing advantage for light coloured females does not exist at high temperatures. Dark individuals suffer lower predation rates at high altitude than do light individuals; the opposite is not true at low elevation. Duration of diel activity is markedly extended for darker butterflies under cool conditions, but only slightly for light individuals during warm sunny periods. Melanistic individuals are able to prolong the duration of activity for feeding, mate location, oviposition and escape from predation under cold conditions. This appears to the prime benefit for melanism in this population. This is the first demonstration, in a natural population, of the benefit of alpine melanism in insects. / Science, Faculty of / Zoology, Department of / Graduate

The grey squirrel (Sciurus carolinensis) has wildtype and melanic (dark) colour morphs. Melanism is associated with variations in the melanocortin-1 receptor (MC1R) gene in a number of species. The MC1R protein is a G-protein coupled receptor, predominantly expressed in melanocytes, where it is a key regulator of pigment production. To investigate the genetic and molecular basis of melanism, the MC1R genes of the wildtype and melanic grey squirrel were sequenced. The wildtype (MC1R-wt) and melanic (MC1RΔ24) variants of the MC1R were then functionally characterised in a cell-based assay. The MC1R gene of the grey squirrel was found to have a 24 base pair (bp) deletion associated with melanism. The MC1R is typically activated by its agonist, the alpha-melanocyte stimulating hormone (α-MSH), which stimulates dark pigment production by raising intracellular cAMP levels. Conversely, the MC1R is inactivated by its inverse agonist, the agouti signalling protein (ASIP), which stops dark pigment production by lowering intracellular cAMP levels. To investigate the effects that the 24 bp deletion have on receptor function, MC1R-wt and MC1RΔ24 genes were transfected into HEK293 cells. Cells expressing either MC1R-wt or MC1RΔ24 were stimulated with α-MSH or ASIP and intracellular cAMP levels were measured. Unstimulated MC1RΔ24 cells showed higher basal activity than the MC1R-wt cells. Both MC1R-wt and MC1RΔ24 cells responded to α-MSH with a concentration-dependent increase in intracellular cAMP. However, while the MC1Rwt cells responded to ASIP with a concentration-dependent decrease in intracellular cAMP, MC1RΔ24 cells responded with an increase in cAMP. Melanism in the grey squirrel is associated with a 24 bp deletion in the MC1R. Cells expressing MC1RΔ24 have higher basal levels of cAMP than MC1R-wt cells. ASIP acts as an inverse agonist to the MC1R-wt but as an agonist to the MC1RΔ24. As MC1RΔ24 cells have higher levels of cAMP, and higher levels of cAMP lead to dark pigment production, the 24 bp deletion is the likely molecular cause of melanism in the grey squirrel.

Thesis (MSc (Botany and Zoology))--University of Stellenbosch, 1990. / Mouton and Oelofsen (1988) suggested that melanistic cordylid populations in the southwestern Cape represent relict, cold-adapted populations which evolved under adverse climatic conditions during the Last Glacial Period, 18000-16000 BP. The first section of this study was undertaken to test their model for the evolution of melanism in the cordylid species, Cordylus polyzonus. For this purpose, geographic character variation among populations of Cordylus polyzonus in the region south of 32°15' latitude and west of 19015' longitude was investigated. Variation in 122 external morphological characters was analysed in 306 specimens from 93 localities. Although no inter-locality variation was observed in most of the characters, a high degree of concordancy was observed in the geographical variation of three headshield characters. These three headshield characters are considered non-adaptive and can accordingly be regarded as good indicators of genealogical relationship. Since they varied together with other adaptive characters such as melanism and body size, it can be concluded that geographically isolated melanistic populations of Cordylus polyzonus represent relicts of a once larger melanistic population. The data underscore the Mouton-Oelofsen hypothesis for the evolution of melanistic cordylid taxa in the south-western Cape. A further corrolary of the Mouton-Oelofsen hypothesis, is that melanistic cordylids are presently restricted to cool enclaves. This assumption was evaluated in the second section of this study by analysing prevailing climatic conditions associated with the occurrence of extant melanistic populations of the genera Cordylus and Pseudocordylus in the southwestern Cape. Climatic data were obtained from 123 weather stations in the study-area. A close correspondence was found between the distribution of melanistic populations along the westcoast and the primary upwelling zones of the southern Benguela Current. Lower mean daily temperatures and a high incidence of advective sea fog on the adjacent coastal regions are direct effects of these cold upwelled waters. Likewise, montane melanistic populations, occurring at relatively high altitudes along the extreme western borders of the Cape Fold Mountains, also experience lower mean daily temperatures and a high incidence of orographic fog and cloud cover. Melanistic cordylid populations therefore generally have to 4. contend with relatively exposed environmental conditions of low temperature and limited solar radiation due to the filtering effect of fog and cloud cover. The fact that all the melanistic taxa in this region occur as small isolated populations limited to cool enclaves, suggests that they are presently in a contracted state under strong environmental pressure. It is postulated that palaeoclimatic conditions very similar to climatic conditions presently prevailing in these enclaves were experienced over the entire western coastal region, at least as far as the Orange River, during the Last Glacial Maximum when melanism probably evolved. Our results therefore corroborate the Mouton-Oelofsen hypothesis that isolated melanistic cordylid populations represent cold-adapted relicts. One would accordingly expect melanistic populations to posses some enhanced ability to absorb infrared radiant heat to cope with these conditions of limited sunshine and lower temperatures. In section three of this study, differences in dorsal skin reflectivity between melanistic and turquoise ~. polyzonus populations were quantified. Furthermore, the physiological ability of this species to change body colour was investigated, as well as ontogenetic colour change. Significant differences in dorsal skin reflectivity existed between melanistic and turquoise specimens in the 500-1300 mjl spectral range, demonstrating the greater heat-absorbing capacity of melanistic populations. The data underscore the MoutonOelofsen model that melanistic populations are "cold-adapted" relicts. Furthermore, ontogenetic colour change is a real phenomenon in the melanistic variation of C. polyzonus. In this respect it is unique among the melanistic cordylid taxa in that ontogenetic colour change seemingly does not occur in the other forms. Experiments to investigate short term colour change in response to different temperatures regimes, produced no conclusive results. On the other hand, seasonal colour changes could be demonstrated for both melanistic and turquoise specimens, indicating that this species has the physiological capacity to change colour. It is, however, believed that geographic colour variation in ~. polyzonus cannot be attributed to this capacity, but is rather the result of selection over time. This view is underscored by the results of section one demonstrating that, apart from colour and body size, melanistic and turquoise forms also differ in certain non-adaptive traits; suggesting that they do not belong to the same primary gene pool.

Le changement climatique récent a des conséquences dramatiques pour la biodiversité, à travers des modifications des conditions abiotiques et biotiques. La vulnérabilité d'une espèce au changement climatique peut dépendre de son habitat, de sa position au sein de sa communauté ainsi que de sa physiologie thermique. A cet égard, les espèces ectothermes, dont la température interne dépend directement du milieu extérieur, sont considérées comme particulièrement vulnérables à l'augmentation de température. Nous avons étudié expérimentalement l'impact du réchauffement climatique futur sur une espèce de vertébré ectotherme, le lézard vivipare (Zootoca vivipara). Pour cela nous avons utilisé le Métatron, un système de grands enclos semi-naturels dans lesquels les conditions climatiques peuvent être manipulées. Nous avons étudié l'impact d'un climat futur plus chaud (+2°C) sur la dynamique des populations et leur risque d'extinction, ainsi que sur les capacités d'adaptation des populations par la plasticité phénotypique, la sélection et la dispersion. De plus, nous avons considéré l'impact du changement climatique à l'échelle de la communauté. Nous démontrons que le changement climatique futur a un impact négatif sur les populations de lézard vivipare, avec un risque d'extinction à court terme. Cependant, des moyens d'adaptation existent, à travers des changements de phénologie et de physiologie (mélanisme, préférences thermiques). Enfin, les conséquences du changement climatique ne sont pas limitées à l'impact sur les populations de lézard, mais affectent la communauté toute entière, depuis les communautés de plantes et d'insectes jusqu'aux communautés microbiennes. / Recent global change has dramatic impacts on biodiversity, through modifications in abiotic and biotic factors. Species vulnerability to changing climates depend for instance of its habitat, its position within the community and its thermal physiology. In this respect, ectotherm species are considered particularly vulnerable as their body temperature depend directly on their environment. We experimentally studied the impact of future climate change on an ectotherm vertebrate species, the common lizard (Zootoca vivipara). We used the Metatron, a system of semi-natural enclosures in which climatic conditions can be manipulated. We studied the impact of warmer climatic conditions (+2°C) on common lizard's population dynamics and extinction risk, and on population adaptation capacity through plasticity, selection and dispersal. We further investigated the impact of climate change at the community scale. We demonstrated that future climatic conditions pose a threat to common lizard. However, possibilities of adaptation exist through changes in phenology and physiology (preferred temperature and melanism). Finally, we show that changing climatic conditions have an impact on the entire communities, from plants and insects to microbial communities.