Evolutionary shifts in habitat aridity predict evaporative water loss across squamate reptiles

Aridity is an important determinant of species distributions, shaping both ecological and evolutionary diversity. Lizards and snakes are often abundant in deserts, suggesting a high potential for adaptation or acclimation to arid habitats. However, phylogenetic evidence indicates that squamate diversity in deserts may be more strongly tied to speciation within arid habitats than to convergent evolution following repeated colonization from mesic habitats. To assess the frequency of evolutionary transitions in habitat aridity while simultaneously testing for associated changes in water‐balance physiology, we analyzed estimates of total evaporative water loss (EWL) for 120 squamate species inhabiting arid, semiarid, or mesic habitats. Phylogenetic reconstructions revealed that evolutionary transitions to and from semiarid habitats were much more common than those between arid and mesic extremes. Species from mesic habitats exhibited significantly higher EWL than those from arid habitats, while species from semiarid habitats had intermediate EWL. Phylogenetic comparative methods confirmed this association between habitat aridity and EWL despite phylogenetic signal in each. Thus, the historical colonization of arid habitats by squamates is repeatedly associated with adaptive changes in EWL. This physiological convergence, which may reflect both phenotypic plasticity and genetic adaptation, has likely contributed to the success of squamates in arid environments.     

Growth and mortality in a field population of Amphibolurus lizards exposed to seasonal cold and aridity

The demography of a discrete population of Amphibolurus ornatus lizards in Western Australia was studied over a period of four years. Large differences in the rate of growth of individuals from the same cohort were observed during each year of the study and each animal could be classified as maturing at one, two or three years of age. Fast- and slow growing individuals were shown to suffer differential mortality during seasonal stresses with fast growers being cold hardy and drought sensitive whereas slow-growers were cold sensitive but drought resistant. The potential longevity of fast- and slow-growing animals appears to be identical but the survival of animals in the field depends markedly upon the incidence of climatic extremes, slow growers tending to live longer than fast growers in the study area. Changes that were observed in the size and age structure of the breeding population during the period of study can be interpreted in terms of the weather conditions experienced by the population over this period. The significance of the variability in individual growth rates of Amphibolurus ornutus for the persistence of the study population is discussed.     

栖息地荒漠化对蜥蜴群落组成的影响

栖息地改变往往对动物群落及其物种多样性有重要影响,但是目前关于栖息地荒漠化如何影响蜥蜴的群落组成鲜有研究。在内蒙古自治区鄂尔多斯市准格尔旗和达拉特旗地区通过样线法调查了三类不同荒漠化程度样地上的蜥蜴群落组成差异,用典范对应分析探究了关键的影响因素。结果表明,栖息地荒漠化使蜥蜴的群落组成发生显著变化,从固定沙丘、半固定沙丘到流动沙丘,草原沙蜥均是蜥蜴群落的优势物种,密点麻蜥逐渐减少,丽斑麻蜥则消失于荒漠化严重的半固定沙丘和流动沙丘中。随栖息地荒漠化程度的增加,蜥蜴的数量显著减少,密度显著降低,蜥蜴物种的丰富度、香农-威纳(Shannon-Weaver)多样性指数和皮洛(Pielou)均匀性指数均逐次显著下降,而辛普森(Simpson)优势度指数则逐次显著升高。这说明荒漠化不仅使蜥蜴群落组成变得简单,同时造成了蜥蜴物种多样性的下降甚至丧失。荒漠化也使蜥蜴的栖息条件发生显著变化。植被高度、油蒿比例、裸地比例、隐蔽度、表层土壤含水量和表层土壤孔隙度在三类栖息样地之间均存在极显著差异。典范对应分析结果表明,这些环境因子与蜥蜴群落组成的变化密切相关。总体而言,草原沙蜥偏好裸地,密点麻蜥偏好隐蔽性好且土壤湿润又疏松的区域,而丽斑麻蜥则偏好植被高的区域。     

Ecological species replacement of Liolaemus lizards along a habitat gradient

Summary Ecological species replacement of Liolaemus platei by Liolaemus lemniscatus appears to occur along a north-south oriented herb density gradient. Both congeners are the only ground-dwelling lizards in the area; this suggests that L. lemniscatus is an ecological counterpart of L. platei in herbaceous habitats. Relatively longer legs appear to be associated with the utilization of herb-free habitats by L. platei; the shorter legs possessed by L. lemniscatus are associated with the utilization of habitats of higher herb density. Morphological evidence indicates the presence of intermediate populations in the zone of intermediate herb density. Alternatives of a single-species cline versus interspecific hybridization between the two taxa are discussed.     

Biomass-density relationship varies with water use efficiency across an aridity gradient

Accumulating evidence has shown that the scaling exponent (α) of the aboveground biomass-density relationship is not a constant value. Debate continues over what determines the variation in α. By measuring foliar stable carbon isotope, plant morphological traits and α along an aridity gradient from eastern to western China, we confirmed that the variation in α was accounted for by changes in plant morphological traits which are adaptive strategies for enhancing water use efficiency during drought stress. This information can be crucial for understanding and predicting community and ecosystem processes.     

Diet and habitat use of frillneck lizards in a seasonal tropical environment

A population of frillneck lizards, Chlamydosaurus kingii, was monitored by mark-recapture and telemetry over a 2 year period in Kakadu National Park. The aims of the study were to document changes in diet, growth, condition and habitat use between the wet and dry seasons of northern Australia, in light of recent research examining seasonal variation in the physiology of this species. Frillneck lizards feed on a diverse range of invertebrates in both seasons, even though there is a substantial reduction in food avail-ability in the dry season. Harvester termites from the genus Drepanotermes constitute a major component of the diet, and the prevalence of termites in the diet of sedentary foragers in a tropical environment is unusual. Adult male body condition remained relatively stable throughout the year, but females experienced considerable variation. These differences are attributed to different reproductive roles of the sexes. Growth in C. kingii was restricted to the wet season, when food availability was high, and growth was minimal in the dry season when food availability was low. The method used in catching lizards was an important factor in determining seasonal habitat use. Telemetered lizards selected a significantly different distribution of tree species than was randomly available, and they selected significantly larger tree species during the dry season. Lizards spotted along roadsides showed little seasonal variation in the selection of tree species or tree sizes. The results suggest a comprehensive change in the ecology of this species, in response to an annual cycle of low food and moisture availability, followed by a period with few resource restrictions.     

The relative importance of respiratory water loss in scorpions is correlated with species habitat type and activity pattern

Scorpions exhibit some of the lowest recorded water loss rates compared with those of other terrestrial arthropods of similar body size. Evaporative water loss (EWL) includes cuticular transpiration and respiratory water loss (RWL) from gas exchange surfaces, that is, book lung lamellae. Estimated fractions of cuticular and respiratory losses currently available from the literature show considerable variation, at least partly as a result of differences in methodology. This study reports RWL rates and their relative importance in scorpions from two families (Buthidae and Scorpionidae), including both xeric and mesic species (or subspecies). Two of the included Buthidae were surface-dwelling species, and another inhabits empty burrows of other terrestrial arthropods. This experimental design enabled correlating RWL importance with scorpion phylogeny, habitat type, and/or homing behavior. Buthidae species exhibited significantly lower EWL rates compared with those of Scorpionidae, whereas effects of habitat type and homing behavior were not significant. Resting RWL rates were not significantly affected by scorpion phylogeny, but rates for the xeric species (totaling ~10% of EWL rates at 30°C) were significantly lower compared with those of mesic species. These lower RWL values were correlated with significantly lower H(2)O/CO(2) emission rates in xeric species. The experimental setup and ~24-h duration of each individual recording allowed estimating the effect of interspecific variation in activity on RWL proportions. The high respiratory losses in active hydrated Scorpio maurus fuscus, totaling 30% of EWL, suggest that behavioral discretion in this species is a more likely mechanism for body water conservation under stressful conditions when compared with the responses of other studied species.     

Implications of climate change on the habitat shifts of tropical lizards

The effect of temperature on the distributions of ectothermic vertebrates is well documented. Despite the increase of 6°C expected in the next 60 years in South America, numerous vertebrates are still considered as ‘Least Concern’ species by the IUCN due to their large distribution, insufficient widespread threats and insignificant population decline. One example is the lizard Tropidurus torquatus (Squamata: Tropiduridae), commonly found thermoregulating in anthropic environments throughout the Brazilian Cerrado, but restricted to gallery forests in the equator‐ward localities. The urban areas in this warmer region have been colonised by other closely related congeners (e.g. Tropidurus oreadicus). This study aimed to understand this divergence of habitat selection by these tropirudids that may explain some of the species responses to past and future climate warming. We collected body temperatures (T_b), micro‐environmental temperatures (T_a) and operative (T_e) temperatures in four sites along a latitudinal gradient: a pole‐ward and two central sites where T. torquatus inhabit urban areas and one equator‐ward site where T. torquatus and T. oreadicus occur in the gallery forest and in urban microhabitats, respectively. All three populations of T. torquatus present similar T_b (35.5–36°C) and shared microhabitats with a similar T_a (34–37.3°C). The T_e in the equator‐ward urban site was considerably higher than in the gallery forest. Tropidurus oreadicus T_b was 38.2 °C (30.1–41.3°C) and was active at a T_a of 30.5–42.3°C. The overlap between the genus T_b, T_a and T_e highlights a decrease in the hours of activity that lizards would experience under climate warming. The reduction of hours of activity together with the devastation of natural habitats represents threats and an alarming scenario especially for the equator‐ward populations.     

Relationships among morphology, clinging performance and habitat use in Liolaemini lizards

The central tenet of ecomorphological theory holds that different ecological requirements lead to different organismal designs (morphology). Here, we studied the relationships between performance (interlocking grasping) and forelimb morphological traits in species of lizards that exploit different structural habitats in a phylogenetic context. The performance (measured by the maximum force of clinging to substrate) was measured on different substrate types. After phylogenetically informed analyses, we found that arboreal and saxicolous species showed stronger resistance to mechanical traction in all substrates when compared to generalists and sand dweller lizards. These species showed a positive relationship between forelimb dimensions (humerus length and length of claw of toe 5) and maximum force exerted, on the contrary, hand width, claw height (CH) of digits III and IV and claw length of toe 4 showed a negative relationship. In addition, we observed a partial positive correlation between CH and maximal cling force on rough surfaces, but not on smooth surfaces.     

Adaptation of metabolism and evaporative water loss along an aridity gradient

Broad-scale comparisons of birds indicate the possibility of adaptive modification of basal metabolic rate (BMR) and total evaporative water loss (TEWL) in species from desert environments, but these might be confounded by phylogeny or phenotypic plasticity. This study relates variation in avian BMR and TEWL to a continuously varying measure of environment, aridity. We test the hypotheses that BMR and TEWL are reduced along an aridity gradient within the lark family (Alaudidae), and investigate the role of phylogenetic inertia. For 12 species of lark, BMR and TEWL decreased along a gradient of increasing aridity, a finding consistent with our proposals. We constructed a phylogeny for 22 species of lark based on sequences of two mitochondrial genes, and investigated whether phylogenetic affinity played a part in the correlation of phenotype and environment. A test for serial independence of the data for mass-corrected TEWL and aridity showed no influence of phylogeny on our findings. However, we did discover a significant phylogenetic effect in mass-corrected data for BMR, a result attributable to common phylogenetic history or to common ecological factors. A test of the relationship between BMR and aridity using phylogenetic independent constrasts was consistent with our previous analysis: BMR decreased with increasing aridity.     

Experimental manipulation reveals the importance of refuge habitat temperature selected by lizards

Refuges provide shelter from predators, and protection from exposure to the elements, as well as other fitness benefits to animals that use them. In ectotherms, thermal benefits may be a critical aspect of refuges. We investigated microhabitat characteristics of refuges selected by a heliothermic scincid lizard, Carlia rubrigularis, which uses rainforest edges as habitat. We approached lizards in the field, simulating a predator attack, and quantified the refuge type used, and effect of environmental temperatures (air temperature, substrate temperature and refuge substrate temperature) on the amount of time skinks remained in refuges after hiding (emergence time). In respone to our approach, lizards were most likely to flee into leaf litter, rather than into rocks or woody debris, and emergence time was dependent on refuge substrate temperature, and on refuge substrate temperature relative to substrate temperature outside the refuge. Lizards remained for longer periods in warmer refuges, and in refuges that were similar in temperature to outside. We examined lizard refuge choice in response to temperature and substrate type in large, semi‐natural outdoor enclosures. We experimentally manipulated refuge habitat temperature available to lizards, and offered them equal areas of leaf litter, woody debris and rocks. When refuge habitat temperature was unmanipulated, lizards (85%) preferred leaf litter, as they did in the field. However, when we experimentally manipulated the temperature of the leaf litter by shading, most skinks (75%) changed their preferred refuge habitat from leaf litter to woody debris or rocks. These results suggest that temperature is a critical determinant of refuge habitat choice for these diurnal ectotherms, both when fleeing from predators and when selecting daytime retreats.     

Summer annual plants exhibit germination responses to an aridity gradient consistent with both delayed germination and phenotypic plasticity

Plant Ecology - Two adaptive strategies to face variable environments have been proposed for annual plants in deserts: delayed and plastic (predictive) germination. All theories and models have...     

The visual ecology of Puerto Rican anoline lizards: habitat light and spectral sensitivity

The visual ecology of six closely related species of Puerto Rican anoline lizards was investigated and they were found to occupy four distinct habitat types in terms of light conditions: “full shade”, “partial shade”, “no shade”, and “forest canopy.”The habitats differed substantially in total radiance and irradiance as well as in the shape of the irradiance spectrum. The shape of the radiance spectrum was similar in all of the habitats. We used electroretinogram (ERG) flicker photometry to measure spectral sensitivity and found the curves for all six species to be similar. The spectral sensitivity peaked in the range 550–560 nm, which matched the peak in spectral radiance for all of the habitats. The shape of the spectral-sensitivity curve was similar to those of a number of other terrestrial vertebrates. We suggest that the convergence of the shape of the photopic ERG-determined spectral-sensitivity curve in many terrestrial vertebrates may, in part, be due to the fact that the background radiance of many terrestrial habitats is dominated by the reflectance spectrum of green vegetation which peaks at 550 nm. Accepted: 14 May 1997     

Energy metabolism and evaporative water loss of Pristidactylus lizards

The effects of thermal conditions on the components of the physiological traits, energy metabolism and evaporative water loss, were studied in two species of Pristidactylus lizards, from different thermal habitats, forest and scrubland areas. The compared species had similar average daily metabolic rates, pulmo-cutaneous and pulmonary evaporative water loss. They differ in the daily distribution of energy expenditure, and in the voluntary aerobic scope. Similarities in most of the studied variables, may reflect either functional or historical constraints on these variables.     

Phylogenetic evidence for multiple losses of a sexually selected character in phrynosomatid lizards

The evolution of conspicuous male display ornaments is a common trend in diverse groups of organisms and a continuing challenge to studies of sexual selection. A phylogenetic approach was used to examine macro-evolutionary patterns of change in sexually dichromatic display coloration (distinctively coloured belly patches) among 130 taxa of phrynosomatid lizards. The results showed repeated losses of sexual dimorphism, which occur through losses of conspicuous male coloration or gains of conspicuous female coloration. The frequent loss of male traits is surprising, given that sexual selection presumably drives their evolutionary origin and maintenance, but is consistent with a recently proposed hypothesis suggesting that females may lose responsiveness to male traits over macro-evolutionary time-scales. The observation of repeated losses of male traits in phrynosomatid lizards (and other groups) may have implications for testing among competing models for the evolution of female preferences. A concentrated changes test showed that changes in male display coloration are significantly associated with the use of ground-dwelling habitat, as opposed to rock- or tree-dwelling habitats. This result suggests a role for natural selection in the loss of male display traits in phrynosomatid lizards, but habitat type alone may be insufficient to explain these losses.     

The relationship between sexual size dimorphism and habitat use in Greater Antillean Anolis lizards

Abstract.— Sexual size dimorphism (SSD) is the evolutionary result of selection operating differently on the body sizes of males and females. Anolis lizard species of the Greater Antilles have been classified into ecomorph classes, largely on the basis of their structural habitat (perch height and diameter). We show that the major ecomorph classes differ in degree of SSD. At least two SSD classes are supported: high SSD (trunk-crown, trunk-ground) and low SSD (trunk, crown-giant, grass-bush, twig). Differences cannot be attributed to an allometric increase of SSD with body size or to a phylogenetic effect. A third explanation, that selective pressures on male and/or female body size vary among habitat types, is examined by evaluating expectations from the major relevant kinds of selective pressures. Although no one kind of selective pressure produces expectations consistent with all of the information, competition with respect to structural habitat and sexual selection pressures are more likely possibilities than competition with respect to prey size or optimal feeding pressures. The existence of habitat-specific sexual dimorphism suggests that adaptation of Anolis species to their environment is more complex than previously appreciated.     

The relevance of morphology for habitat use and locomotion in two species of wall lizards

Understanding if morphological differences between organisms that occupy different environments are associated to differences in functional performance can suggest a functional link between environmental and morphological variation. In this study we examined three components of the ecomorphological paradigm – morphology, locomotor performance and habitat use – using two syntopic wall lizards endemic to the Iberian Peninsula as a case study to establish whether morphological variation is associated with habitat use and determine the potential relevance of locomotor performance for such an association. Differences in habitat use between both lizards matched patterns of morphological variation. Indeed, individuals of Podarcis guadarramae lusitanicus, which are more flattened, used more rocky environments, whereas Podarcis bocagei, which have higher heads, used more vegetation than rocks. These patterns translated into a significant association between morphology and habitat use. Nevertheless, the two species were only differentiated in some of the functional traits quantified, and locomotor performance did not exhibit an association with morphological traits. Our results suggest that the link between morphology and habitat use is mediated by refuge use, rather than locomotor performance, in this system, and advise caution when extrapolating morphology-performance-environment associations across organisms.