Provenance matters: thermal reaction norms for embryo survival among sockeye salmon Oncorhynchus nerka populations

Differences in thermal tolerance during embryonic development in Fraser River sockeye salmon Oncorhynchus nerka were examined among nine populations in a controlled common‐garden incubation experiment. Forcing embryonic development at an extreme temperature (relative to current values) of 16° C, representing a future climate change scenario, significantly reduced survival compared to the more ecologically moderate temperature of 10° C (55% v. 93%). Survival at 14° C was intermediate between the other two temperatures (85%). More importantly, this survival response varied by provenance within and between temperature treatments. Thermal reaction norms showed an interacting response of genotype and environment (temperature), suggesting that populations of O. nerka may have adapted differentially to elevated temperatures during incubation and early development. Moreover, populations that historically experience warmer incubation temperatures at early development displayed a higher tolerance for warm temperatures. In contrast, thermal tolerance does not appear to transcend life stages as adult migration temperatures were not related to embryo thermal tolerance. The intra‐population variation implies potential for thermal tolerance at the species level. The differential inter‐population variation in thermal tolerance that was observed suggests, however, limited adaptive potential to thermal shifts for some populations. This infers that the intergenerational effects of increasing water temperatures may affect populations differentially, and that such thermally mediated adaptive selection may drive population, and therefore species, persistence.     

Male sterility at extreme temperatures: a significant but neglected phenomenon for understanding Drosophila climatic adaptations

The thermal range for viability is quite variable among Drosophila species and it has long been known that these variations are correlated with geographic distribution: temperate species are on average more cold tolerant but more heat sensitive than tropical species. At both ends of their viability range, sterile males have been observed in all species investigated so far. This symmetrical phenomenon restricts the temperature limits within which permanent cultures can be kept in the laboratory. Thermal heat sterility thresholds are very variable across species from 23 degrees C in heat sensitive species up to 31 degrees C in heat tolerant species. In Drosophila melanogaster, genetic variations are observed among geographic populations. Tropical populations are more tolerant to heat induced sterility and recover more rapidly than temperate ones. A genetic analysis revealed that about 50% of the difference observed between natural populations was due to the Y chromosome. Natural populations have not reached a selection limit, however: thermal tolerance was still increased by keeping strains at a high temperature, close to the sterility threshold. On the low temperature side, a symmetrical reverse phenomenon seems to exist: temperate populations are more tolerant to cold than tropical ones. Compared to Mammals, drosophilids exhibit two major differences: first, male sterility occurs not only at high temperature, but also at a low temperature; second, sterility thresholds are not evolutionarily constrained, but highly variable. Altogether, significant and sometimes major genetic variations have been observed between species, between geographic races of the same species, and even between strains kept in the laboratory under different thermal regimes. In each case, it is easily argued that the observed variations correspond to adaptations to climatic conditions, and that male sterility is a significant component of fitness and a target of natural selection.     

The sea urchin Lytechinus variegatus lives close to the upper thermal limit for early development in a tropical lagoon

Thermal tolerance shapes organisms' physiological performance and limits their biogeographic ranges. Tropical terrestrial organisms are thought to live very near their upper thermal tolerance limits, and such small thermal safety factors put them at risk from global warming. However, little is known about the thermal tolerances of tropical marine invertebrates, how they vary across different life stages, and how these limits relate to environmental conditions. We tested the tolerance to acute heat stress of five life stages of the tropical sea urchin Lytechinus variegatus collected in the Bahía Almirante, Bocas del Toro, Panama. We also investigated the impact of chronic heat stress on larval development. Fertilization, cleavage, morula development, and 4‐armed larvae tolerated 2‐h exposures to elevated temperatures between 28–32°C. Average critical temperatures (LT₅₀) were lower for initiation of cleavage (33.5°C) and development to morula (32.5°C) than they were for fertilization (34.4°C) or for 4‐armed larvae (34.1°C). LT₅₀ was even higher (34.8°C) for adults exposed to similar acute thermal stress, suggesting that thermal limits measured for adults may not be directly applied to the whole life history. During chronic exposure, larvae had significantly lower survival and reduced growth when reared at temperatures above 30.5°C and did not survive chronic exposures at or above 32.3°C. Environmental monitoring at and near our collection site shows that L. variegatus may already experience temperatures at which larval growth and survival are reduced during the warmest months of the year. A published local climate model further suggests that such damaging warm temperatures will be reached throughout the Bahía Almirante by 2084. Our results highlight that tropical marine invertebrates likely have small thermal safety factors during some stages in their life cycles, and that shallow‐water populations are at particular risk of near future warming.     

Thermal tolerance ranges and climate variability: A comparison between bivalves from differing climates

The climate variability hypothesis proposes that in variable temperate climates poikilothermic animals have wide thermal tolerance windows, whereas in constant tropical climates they have small thermal tolerance windows. In this study we quantified and compared the upper and lower lethal thermal tolerance limits of numerous bivalve species from a tropical (Roebuck Bay, north western Australia) and a temperate (Wadden Sea, north western Europe) tidal flat. Species from tropical Roebuck Bay had higher upper and lower lethal thermal limits than species from the temperate Wadden Sea, and Wadden Sea species showed an ability to survive freezing temperatures. The increased freezing resistance of the Wadden Sea species resulted in thermal tolerance windows that were on average 7 °C greater than the Roebuck Bay species. Furthermore, at a local-scale, the upper lethal thermal limits of the Wadden Sea species were positively related to submersion time and thus to encountered temperature variation, but this was not the case for the Roebuck Bay species. A review of previous studies, at a global scale, showed that upper lethal thermal limits of tropical species are closer to maximum habitat temperatures than the upper lethal thermal limits of temperate species, suggesting that temperate species are better adapted to temperature variation. In this study, we show for the first time, at both local and global scales, that the lethal thermal limits of bivalves support the climate variability effect in the marine environment.     

Correlated evolution of thermal niches and functional physiology in tropical freshwater fishes

The role of ecology in phenotypic and species diversification is widely documented. Nonetheless, numerous nonadaptive processes can shape realized niches and phenotypic variation in natural populations, complicating inferences about adaptive evolution at macroevolutionary scales. We tested for evolved differences in thermal tolerances and their association with the realized thermal niche (including metrics describing diurnal and seasonal patterns of temperature extremes and variability) across a genus of tropical freshwater fishes reared in a standardized environment. There was limited evolution along the thermal niche axis associated with variation in maximum temperature and in upper thermal limits. In contrast, there was considerable diversification along the first major axis of the thermal niche associated with minimum temperatures and in lower thermal limits. Across our adaptive landscape analyses, 70% of species exhibited evidence of divergence in thermal niches. Most importantly, the first two major axes of thermal niche variation were significantly correlated with variation in lower thermal limits. Our results indicate adaptation to divergent thermal niches and adaptive evolution of related functional traits, and highlight the importance of divergence in lower thermal limits for the evolution of tropical biodiversity.     

A comparative analysis of the upper thermal tolerance limits of eastern Pacific porcelain crabs, genus Petrolisthes: influences of latitude, vertical zonation, acclimation, and phylogeny

Marine intertidal organisms are subjected to a variety of abiotic stresses, including aerial exposure and wide ranges of temperature. Intertidal species generally have higher thermal tolerance limits than do subtidal species, and tropical species have higher thermal tolerance limits than do temperate species. The adaptive significance of upper thermal tolerance limits of intertidal organisms, however, has not been examined within a comparative context. Here, we present a comparative analysis of the adaptive significance of upper thermal tolerance limits in 20 congeneric species of porcelain crabs, genus Petrolisthes, from intertidal and subtidal habitats throughout the eastern Pacific. Upper thermal tolerance limits are positively correlated with surface water temperatures and with maximal microhabitat temperatures. Analysis of phylogenetically independent contrasts (from a phylogenetic tree on the basis of the 16s rDNA gene sequence) suggests that upper thermal tolerance limits have evolved in response to maximal microhabitat temperatures. Upper thermal tolerance limits increased during thermal acclimation at elevated temperatures, the amount of increase being greater for subtidal than for intertidal species. This result suggests that the upper thermal tolerance limits of some intertidal species may be near current habitat temperature maxima, and global warming thus may affect the distribution limits of intertidal species to a greater extent than for subtidal species.     

Adaptive divergence in embryonic thermal plasticity among Atlantic salmon populations

In the context of global changes, the long‐term viability of populations of endangered ectotherms may depend on their adaptive potential and ability to cope with temperature variations. We measured responses of Atlantic salmon embryos from four populations to temperature variations and used a Q_ST–F_ST approach to study the adaptive divergence among these populations. Embryos were reared under two experimental conditions: a low temperature regime at 4 °C until eyed‐stage and 10 °C until the end of embryonic development and a high temperature regime with a constant temperature of 10 °C throughout embryonic development. Significant variations among populations and population × temperature interactions were observed for embryo survival, incubation time and length. Q_ST was higher than F_ST in all but one comparison suggesting an important effect of divergent selection. Q_ST was also higher under the high‐temperature treatment than at low temperature for length and survival due to a higher variance among populations under the stressful warmer treatment. Interestingly, heritability was lower for survival under high temperature in relation to a lower additive genetic variance under that treatment. Overall, these results reveal an adaptive divergence in thermal plasticity in embryonic life stages of Atlantic salmon suggesting that salmon populations may differentially respond to temperature variations induced by climate change. These results also suggest that changes in temperature may alter not only the adaptive potential of natural populations but also the selection regimes among them.     

Multivariate analysis of adaptive capacity for upper thermal limits in Drosophila simulans

Thermal tolerance is an important factor influencing the distribution of ectotherms, but our understanding of the ability of species to evolve different thermal limits is limited. Based on univariate measures of adaptive capacity, it has recently been suggested that species may have limited evolutionary potential to extend their upper thermal limits under ramping temperature conditions that better reflect heat stress in nature. To test these findings more broadly, we used a paternal half‐sibling breeding design to estimate the multivariate evolutionary potential for upper thermal limits in Drosophila simulans. We assessed heat tolerance using static (basal and hardened) and ramping assays. Our analyses revealed significant evolutionary potential for all three measures of heat tolerance. Additive genetic variances were significantly different from zero for all three traits. Our G matrix analysis revealed that all three traits would contribute to a response to selection for increased heat tolerance. Significant additive genetic covariances and additive genetic correlations between static basal and hardened heat‐knockdown time, marginally nonsignificant between static basal and ramping heat‐knockdown time, indicate that direct and correlated responses to selection for increased upper thermal limits are possible. Thus, combinations of all three traits will contribute to the evolution of upper thermal limits in response to selection imposed by a warming climate. Reliance on univariate estimates of evolutionary potential may not provide accurate insight into the ability of organisms to evolve upper thermal limits in nature.     

Limited potential for adaptation to climate change in a broadly distributed marine crustacean

The extent to which acclimation and genetic adaptation might buffer natural populations against climate change is largely unknown. Most models predicting biological responses to environmental change assume that species' climatic envelopes are homogeneous both in space and time. Although recent discussions have questioned this assumption, few empirical studies have characterized intraspecific patterns of genetic variation in traits directly related to environmental tolerance limits. We test the extent of such variation in the broadly distributed tidepool copepod Tigriopus californicus using laboratory rearing and selection experiments to quantify thermal tolerance and scope for adaptation in eight populations spanning more than 17° of latitude. Tigriopus californicus exhibit striking local adaptation to temperature, with less than 1 per cent of the total quantitative variance for thermal tolerance partitioned within populations. Moreover, heat-tolerant phenotypes observed in low-latitude populations cannot be achieved in high-latitude populations, either through acclimation or 10 generations of strong selection. Finally, in four populations there was no increase in thermal tolerance between generations 5 and 10 of selection, suggesting that standing variation had already been depleted. Thus, plasticity and adaptation appear to have limited capacity to buffer these isolated populations against further increases in temperature. Our results suggest that models assuming a uniform climatic envelope may greatly underestimate extinction risk in species with strong local adaptation.     

GEOGRAPHIC VARIATION IN THERMAL TRAITS IN DIGENEA SIMPLEX AND CHAMPIA PARVULA (RHODOPHYTA) IN RELATION TO PRESENT AND GLACIAL TEMPERATURE REGIMES

Geographic variation in temperature responses (survival and growth) was investigated in two red algae: Digenea simplex (Wulfen) C. Agardh and Champia parvula (C. Agardh) Harvey. D. simplex has a tropical to warm temperate distribution; C. parvula extends from the tropics into the cold temperate zone. Ecoclinal variation was found in both species but was much stronger in C. parvula than in D. simplex. The former species showed variation in upper and lower tolerance limits as well as in the upper and lower limits for growth. The latter species showed variation mainly in its lower tolerance limit. Ecoclinal variation was related to the amount of present and glacial selection pressure along the climate gradient. In both species, isolates from the colder localities had insufficient cold tolerance to have survived low glacial winter temperatures, so these locations must have been colonized after the end of the glaciation. Eastern Mediterranean and Atlantic populations were probably isolated during the glaciation by a thermal barrier at the entrance of the Mediterranean. In C. parvula, evidence existed for a trade-off between the performance at high and at low temperatures, which would enhance selection pressure in opposite directions at either end of the climatic range. No evidence for such a trade-off was found in D. simplex.     

Increases in the evolutionary potential of upper thermal limits under warmer temperatures in two rainforest Drosophila species

Tropical and subtropical species represent the majority of biodiversity. These species are predicted to lack the capacity to evolve higher thermal limits in response to selection imposed by climatic change. However, these assessments have relied on indirect estimates of adaptive capacity, using conditions that do not reflect environmental changes projected under climate change. Using a paternal half‐sib full‐sib breeding design, we estimated the additive genetic variance and narrow‐sense heritability for adult upper thermal limits in two rainforest‐restricted species of Drosophila reared under two thermal regimes, reflecting increases in seasonal temperature projected for the Wet Tropics of Australia and under standard laboratory conditions (constant 25°C). Estimates of additive genetic variation and narrow‐sense heritability for adult heat tolerance were significantly different from zero in both species under projected summer, but not winter or constant, thermal regimes. In contrast, significant broad‐sense genetic variation was apparent in all thermal regimes for egg‐to‐adult viability. Environment‐dependent changes in the expression of genetic variation for adult upper thermal limits suggest that predicting adaptive responses to climate change will be difficult. Estimating adaptive capacity under conditions that do not reflect future environmental conditions may provide limited insight into evolutionary responses to climate change.     

Evolution of thermotolerance in seasonal environments: the effects of annual temperature variation and life-history timing in Wyeomyia smithii

In organisms with complex life cycles, the adaptive value of thermotolerance depends on life-history timing and seasonal temperature profiles. We illustrate this concept by examining variation in annual thermal environments and thermal acclimation among four geographic populations of the pitcher plant mosquito. Only diapausing larvae experience winter, whereas both postdiapause and nondiapause adults occur only during the growing season. Thus, adults experience transient cold stress primarily during the spring. We show that adult cold tolerance (chill coma recovery) is enhanced in spring-like conditions via thermal acclimation but is unaffected by diapause state. Moreover, adult mosquitoes from northern populations were more cold tolerant than those from southern populations largely because acclimation responses were steeper in the north. In contrast to cold tolerance, there was no significant acclimation of heat tolerance (heat knockdown), and no significant differences in heat tolerance between northern and southern populations. Field temperature data show that because of evolved differences in diapause timing, adult exposure to cold stress is remarkably consistent across geography. This suggests that geographic variation in cold tolerance may not be the result of direct selection on adults. Our results illustrate the importance of the interplay between phenological and thermal adaptation for understanding variation along climatic gradients.     

No patterns in thermal plasticity along a latitudinal gradient in Drosophila simulans from eastern Australia

Phenotypic plasticity may be an important initial mechanism to counter environmental change, yet we know relatively little about the evolution of plasticity in nature. Species with widespread distributions are expected to have evolved higher levels of plasticity compared with those with more restricted, tropical distributions. At the intraspecific level, temperate populations are expected to have evolved higher levels of plasticity than their tropical counterparts. However, empirical support for these expectations is limited. In addition, no studies have comprehensively examined the evolution of thermal plasticity across life stages. Using populations of Drosophila simulans collected from a latitudinal cline spanning the entire east coast of Australia, we assessed thermal plasticity, measured as hardening capacity (the difference between basal and hardened thermal tolerance) for multiple measures of heat and cold tolerance across both adult and larval stages of development. This allowed us to explicitly ask whether the evolution of thermal plasticity is favoured in more variable, temperate environments. We found no relationship between thermal plasticity and latitude, providing little support for the hypothesis that temperate populations have evolved higher levels of thermal plasticity than their tropical counterparts. With the exception of adult heat survival, we also found no association between plasticity and ten climatic variables, indicating that the evolution of thermal plasticity is not easily predicted by the type of environment that a particular population occupies. We discuss these results in the context of the role of plasticity in a warming climate.     

极端生理耐受性导致果蝇Zaprionus indianus 在温带地区广泛分布（英文）

在过去的十年中, Zaprionus indianus这一温带适应性果蝇已经入侵印度次大陆, 并扩大了其在该地区的分布。Z. indianus能成功入侵是由于它具有很强的适应性和对极端生理条件的耐受性。对Z. indianus (温带狭域分布种类) 和黑腹果蝇Drosophila melanogaster (全球广域分布种类)在极端温度下未成熟期和成虫期发育阈值的比较研究表明, 两者的死亡率和发育起点温度存在显著差异。为了检测越冬期间未成熟期和成虫期抗逆性和存活率的变化, 以采自印度温带和热带不同地点的Z. indianus种群进行饲养实验。在温带地区的田间养虫笼中以及恒定的实验室条件下监测这些种群的卵孵化率和成虫存活率, 直至全部成虫死亡。结果表明, 由于温带地区卵孵化率和存活率高, 导致总的孵化率和存活率在不同纬度间存在显著差异。卵至成虫发育实验结果表明, 低温条件下产下的卵在温度适中时成功发育成成虫。由此可见, 这种昆虫在未成熟期具有的气候适应性以及在成虫期具有的抗逆性可为该物种提供季节性保护。考虑到气候变暖情况, 即温度增加0.6℃, 温度的少许改变都可能导致种群存活能力的显著增强和发育历期缩短。这些结果可解释Z. indianus为什么能够轻易突破障碍并适应新的环境。     

Evidence for a freezing tolerance-growth rate trade-off in the live oaks (Quercus series Virentes) across the tropical-temperate divide

? It has long been hypothesized that species are limited to the north by minimum temperature and to the south by competition, resulting in a trade-off between freezing tolerance and growth rate. We investigated the extent to which the climatic origins of populations from four live oak species (Quercus series Virentes) were associated with freezing tolerance and growth rate, and whether species fitted a model of locally adapted populations, each with narrow climatic tolerances, or of broadly adapted populations with wide climatic tolerances. ? Acorns from populations of four species across a tropical-temperate gradient were grown under common tropical and temperate conditions. Growth rate, seed mass, and leaf and stem freezing traits were compared with source minimum temperatures. ? Maximum growth rates under tropical conditions were negatively correlated with freezing tolerance under temperate conditions. The minimum source temperature predicted the freezing tolerance of populations under temperate conditions. The tropical species Q. oleoides was differentiated from the three temperate species, and variation among species was greater than among populations. ? The trade-off between freezing tolerance and growth rate supports the range limit hypothesis. Limited variation within species indicates that the distributions of species may be driven more strongly by broad climatic factors than by highly local conditions.     

Differences in egg thermotolerance between tropical and temperate populations of the migratory locust Locusta migratoria (Orthoptera: Acridiidae)

The migratory locust Locusta migratoria L., which is widely distributed throughout the world, exhibits within- and between-population variation in cold tolerance. To understand physiological adaptation in populations, we studied the genetic basis of thermotolerance in Hainan (tropical) and Liaoning (temperate) populations and measured expression of Hsp70 and Hsp90 mRNA in both populations at low (0 degrees C) and high temperatures (40 degrees C). Phenotypic variation of thermotolerance is heritable. Heritable characteristics differed among different stages of locust egg development, as well as among different measures of thermotolerance. Nuclear genetic factors, rather than cytoplasmic factors, contribute to differences in cold tolerance between the tropical and temperate populations of the migratory locust; for heat tolerance, maternal effects were involved in three stages of egg development. Expression of Hsp90 mRNA was induced in temperate population after heat shock (40 degrees C x 12h), whereas expression of Hsp70 and 90 was induced in tropical population after cold shock (0 degrees C x 12h). We suggest that thermotolerance of locust eggs has a complex genetic basis and heat shock proteins may be involved in differences of thermotolerance between locust populations.     

Vulnerability to climate warming and acclimation capacity of tropical and temperate coastal organisms

Ecological forecasting on the likely impacts of climate warming is crucial at a time when several ecosystems seem to be responding to this environmental threat. Among the most important questions are: which are the most vulnerable organisms to climate warming and where are they? Recently, there has been debate on whether the tropics or temperate zones are more vulnerable to warming. Vulnerability toward higher temperatures will depend on the organisms’ thermal limits and also on their acclimation capacity, which remains largely unknown for most species. The aim of the present work was to estimate (1) the upper thermal limits (Critical Thermal Maximum (CTMax)), (2) the warming tolerance (CTMax – Maximum Habitat Temperature) and (3) the acclimation capacity of tropical and temperate rocky shore organisms. Differences in biological groups (decapod crustaceans vs fish) were investigated and the effect of region (tropical vs temperate) and habitat (intertidal vs subtidal) was tested. Overall, 35 species were tested. For the assessment of the acclimation capacity, tropical-temperate pairs of closely related species of shrimp, crab and fish were selected. Warming tolerance was higher for temperate species than for tropical species and higher for subtidal species than for intertidal species, confirming that species with the highest thermal limits have the lowest warming tolerance. All species tested presented some acclimation capacity (CTMax_Trial − CTMax_Control), with the exception of gobiid fish, which was not observed to acclimate. The tropical species tested showed a lower acclimation capacity than their temperate counterparts. Given that tropical rocky shore organisms are already living very close to their thermal limits and that their acclimation capacity is limited, it is likely that the impacts of global warming will be evident sooner in the tropics than in the temperate zone.     

CTmax in brook trout (Salvelinus fontinalis) embryos shows an acclimation response to developmental temperatures but is more variable than in later life stages

Critical thermal maximum (CT_max) is widely used to measure upper thermal tolerance in fish but is rarely examined in embryos. Upper thermal limits generally depend on an individual's thermal history, which molds plasticity. We examined how thermal acclimation affects thermal tolerance of brook trout (Salvelinus fontinalis) embryos using a novel method to assess CT_max in embryos incubated under three thermal regimes. Warm acclimation was associated with an increase in embryonic upper thermal tolerance. However, CT_max variability was markedly higher than is typical for juvenile or adult salmonids.     

Effect of Body Size on Reef Fish Tolerance to Extreme Low and High Temperatures

Understanding the thermal physiology of tropical marine organisms has become an issue of major interest due to the potential impact of current global changes in temperature. In this study we report the effect of body size on the thermal tolerance (as critical thermal maximum (CTMax) and minimum (CTMin)) of seven reef fish species from Gorgona Island (tropical eastern Pacific Ocean). Within the studied species we found little variation in CTMax and CTMin among fishes ranging from juveniles to adults. This suggests that thermal tolerance of small tropical reef fishes is not significantly affected by differences in body size. The reduced intra-specific variation in thermal tolerance found in these species also suggests a limited capability to adapt to extreme thermal conditions and raises concerns regarding current global changes in temperature.     

Desiccation tolerance and starvation resistance exhibit opposite latitudinal clines in Indian geographical populations of Drosophila kikkawai

Abstract. 1. Desiccation tolerance and starvation resistance demonstrated significant differentiation among seven Indian geographical populations of Drosophila kikkawai, collected along a latitudinal range of 12.6–32.7 °N. Lack of significant differences in two successive generations suggested that these physiological traits were genetically controlled. 2. North Indian populations of D. kikkawai displayed significantly higher desiccation tolerance than southern populations, whereas there was a reverse trend for starvation tolerance (r > 0.90). Regression slope values indicated an increase of 0.61 h for desiccation and a decrease of 1.71 h per degree latitude for starvation tolerance at 17 °C. The traits evidenced opposite latitudinal clines, and such data also matched thermal climatic conditions on the Indian subcontinent. The survival duration for such traits was significantly higher at 17 than at 25 °C. 3. Significantly higher starvation tolerance in south Indian populations might be due to large population size, species interactions, and higher metabolic rates in the humid tropical environments. In contrast, prolonged unfavourable colder climatic conditions are known to favour starvation tolerance in temperate regions. Thus, the causes of desiccation and starvation tolerance seem quite different under tropical and temperate conditions. 4. Starvation tolerance was correlated negatively with body weight and ovariole number, which might be due to a trade-off in favour of greater allocation to non-lipidic reserves for sustaining starvation tolerance in the tropics. Reduction in metabolic rate may not be applicable for observed higher starvation tolerance in the tropical populations. 5. Multiple regression analysis demonstrated a major effect of coefficient of variation of mean monthly temperature for both the traits of ecological significance. Thus, Indian geographical populations of D. kikkawai provided evidence of independent genetic divergence for starvation and desiccation tolerance under natural conditions.