The roles of genetic drift and natural selection in quantitative trait divergence along an altitudinal gradient in Arabidopsis thaliana

Understanding how natural selection and genetic drift shape biological variation is a central topic in biology, yet our understanding of the agents of natural selection and their target traits is limited. We investigated to what extent selection along an altitudinal gradient or genetic drift contributed to variation in ecologically relevant traits in Arabidopsis thaliana. We collected seeds from 8 to 14 individuals from each of 14 A. thaliana populations originating from sites between 800 and 2700 m above sea level in the Swiss Alps. Seed families were grown with and without vernalization, corresponding to winter-annual and summer-annual life histories, respectively. We analyzed putatively neutral genetic divergence between these populations using 24 simple sequence repeat markers. We measured seven traits related to growth, phenology and leaf morphology that are rarely reported in A. thaliana and performed analyses of altitudinal clines, as well as overall Q_ST-F_ST comparisons and correlation analyses among pair-wise Q_ST, F_ST and altitude of origin differences. Multivariate analyses suggested adaptive differentiation along altitude in the entire suite of traits, particularly when expressed in the summer-annual life history. Of the individual traits, a decrease in rosette leaf number in the vegetative state and an increase in leaf succulence with increasing altitude could be attributed to adaptive divergence. Interestingly, these patterns relate well to common within- and between-species trends of smaller plant size and thicker leaves at high altitude. Our results thus offer exciting possibilities to unravel the underlying mechanisms for these conspicuous trends using the model species A. thaliana.     

Morphological variation and floral abnormalities in a trigger plant across a narrow altitudinal gradient

Local adaptation in alpine plants has been demonstrated across wide altitudinal gradients, but has rarely been examined across the alpine‐to‐montane transition that often encompasses only a few hundred metres. Here we characterize morphological variation in leaf and floral characteristics of the trigger plant Stylidium armeria along a narrow altitudinal gradient in the Bogong High Plains in Victoria. Across this gradient, which encompasses the high‐elevation limit of this species, linear changes were found for floral scape height, leaf length and flower number. All these traits decreased with increasing altitude, whereas the frequency of abnormal flowers increased. When plants were grown in a common garden environment, an altitudinal pattern for flower abnormalities was no longer detected. However, altitudinal patterns for leaf length and scape height were maintained, albeit weaker than in the field. This indicates heritable variation for these morphological traits; the altitudinal patterns are likely to reflect the effects of selection by environmental factors that vary with altitude. Selection pressures remain to be identified but have generated both cogradient and countergradient patterns of variation.     

Leaf morphology shift linked to climate change

Climate change is driving adaptive shifts within species, but research on plants has been focused on phenology. Leaf morphology has demonstrated links with climate and varies within species along climate gradients. We predicted that, given within-species variation along a climate gradient, a morphological shift should have occurred over time due to climate change. We tested this prediction, taking advantage of latitudinal and altitudinal variations within the Adelaide Geosyncline region, South Australia, historical herbarium specimens (n = 255) and field sampling (n = 274). Leaf width in the study taxon, Dodonaea viscosa subsp. angustissima, was negatively correlated with latitude regionally, and leaf area was negatively correlated with altitude locally. Analysis of herbarium specimens revealed a 2 mm decrease in leaf width (total range 1-9 mm) over 127 years across the region. The results are consistent with a morphological response to contemporary climate change. We conclude that leaf width is linked to maximum temperature regionally (latitude gradient) and leaf area to minimum temperature locally (altitude gradient). These data indicate a morphological shift consistent with a direct response to climate change and could inform provenance selection for restoration with further investigation of the genetic basis and adaptive significance of observed variation.     

Selection by Pollinators on Floral Traits in Generalized Trollius ranunculoides (Ranunculaceae) along Altitudinal Gradients

Abundance and visitation of pollinator assemblages tend to decrease with altitude, leading to an increase in pollen limitation. Thus increased competition for pollinators may generate stronger selection on attractive traits of flowers at high elevations and cause floral adaptive evolution. Few studies have related geographically variable selection from pollinators and intraspecific floral differentiation. We investigated the variation of Trollius ranunculoides flowers and its pollinators along an altitudinal gradient on the eastern Qinghai-Tibet Plateau, and measured phenotypic selection by pollinators on floral traits across populations. The results showed significant decline of visitation rate of bees along altitudinal gradients, while flies was unchanged. When fitness is estimated by the visitation rate rather than the seed number per plant, phenotypic selection on the sepal length and width shows a significant correlation between the selection strength and the altitude, with stronger selection at higher altitudes. However, significant decreases in the sepal length and width of T. ranunculoides along the altitudinal gradient did not correspond to stronger selection of pollinators. In contrast to the pollinator visitation, mean annual precipitation negatively affected the sepal length and width, and contributed more to geographical variation in measured floral traits than the visitation rate of pollinators. Therefore, the sepal size may have been influenced by conflicting selection pressures from biotic and abiotic selective agents. This study supports the hypothesis that lower pollinator availability at high altitude can intensify selection on flower attractive traits, but abiotic selection is preventing a response to selection from pollinators.     

Altitudinal and seasonal variation in microsatellite allele frequencies of Drosophila buzzatii

Variation in climate, particularly temperature, is known to affect the genetic composition of populations. Although there have been many studies of latitudinal variation, comparisons of populations across altitudes or seasons, particularly for animal species, are less common. Here, we study genetic variation (microsatellite markers) in populations of Drosophila buzzatii collected along altitudinal gradients and in different seasons. We found no differences in genetic variation between 2 years or between seasons within years. However, there were numerous cases of significant associations between allele frequencies or expected heterozygosities and altitude, with more than half showing nonlinear relationships. While these associations indicate possible selection and local altitudinal adaptation, direct tests gave strong evidence for selection affecting two loci and weaker evidence for five other loci. Two loci that are located within an inversion (including the one with strongest evidence for selection) show a linear increase in genetic diversity with altitude, likely due to thermal selection. Parallel associations with altitude here and with latitude in Australian populations indicate that selection is operating on chromosomal regions marked by some of the loci.     

Are plant pathogen populations adapted for encounter with their host? A case study of phenological synchrony between oak and an obligate fungal parasite along an altitudinal gradient

Biotrophic fungal pathogens are expected to have adapted to their host plants for phenological synchrony, to optimize the possibility of contacts leading to infections. We investigated the patterns and causes of variation in phenological synchrony in the oak‐powdery mildew pathosystem, a major disease in natural ecosystems. The study was carried out along an altitudinal gradient, representing a wide temperature range, in mature oak stands. Both sporulation (pathogen infective stage) and oak flushing (host susceptible stage) were delayed with increasing elevation, but with a significantly different sensitivity for the two species. This resulted in a variable host–pathogen synchrony along the gradient. A common garden experiment did not give evidence of among‐population genetic differentiation (past adaptation) for fungal phenology. This could be explained by the high phenotypic variation in phenology within host populations, precluding selection on fungal phenology at the population scale, but possibly favouring adaptation at the within‐population scale. Phenotypic plasticity was the major cause of the observed variation in the phenology of the fungal populations.     

Effect of altitude on the genetic structure of an Alpine grass, Poa hiemata

Background and Aims

The persistence of plants inhabiting restricted alpine areas under climate change will depend upon many factors including levels of genetic variation in adaptive traits, population structure, and breeding system.

Methods

Using microsatellite markers, the genetic structure of populations of a relatively common alpine grass, Poa hiemata, is examined across three altitudinal gradients within the restricted Australian alpine zone where this species has previously been shown to exhibit local adaptation across a narrow altitudinal gradient.

Key Results

Genetic variation across six microsatellite markers revealed genetic structuring along altitudinal transects, and a reduction in genetic variation at high and low altitude extremes relative to sites central within transects. There was less genetic variation among transect sites compared with altitudinal gradients within transects, even though distances among transects were relatively larger. Central sites within transects were less differentiated than those at extremes.

Conclusions

These patterns suggest higher rates of gene flow among sites at similar altitudes than along transects, a process that could assist altitudinal adaptation. Patterns of spatial autocorrelation and isolation by distance changed with altitude and may reflect altered patterns of dispersal via pollen and/or seed. There was evidence for selfing and clonality in neighbouring plants. Levels of gene flow along transects were insufficient to prevent adaptive changes in morphological traits, given previously measured levels of selection.Key words: Poa hiemata, genetic structure, altitudinal gradient, microsatellite, gene flow, climate change     

Exploring the impact of neutral evolution on intrapopulation genetic differentiation in functional traits in a long-lived plant

Most plant species, particularly long-lived plants, harbor a large amount of genetic variation within populations. A central issue in evolutionary ecology is to explore levels of genetic variation and understand the mechanisms that influence them. In this study, our goals were to examine the impact of neutral evolutionary processes on the genetic variance and functional diversity within three populations of a long-lived plant (Quercus suber L.). For this purpose, we genotyped the progeny of 45 open-pollinated mother trees from three populations originating from Spain, Portugal, and Morocco using six microsatellite markers. Seedlings were planted in a common garden trial and were phenotypically characterized by seven leaf functional traits. Molecular analyses revealed weak genetic differences between Iberian and Moroccan populations. Nevertheless, high genetic differentiation was observed among maternal families within populations. Differentiation between particular maternal families from the same population reached values of 29.2 %, which far exceeds the values reported between the most genetically distant populations for this species (11.7 %). Maternal families differed also in phenology, leaf size, and shape traits. In the Moroccan population, there were correlations among matrices of distances for molecular markers, leaf shape traits (e.g., leaf circularity index), and phenology, indicating that maternal families with contrasting phenologies were genetically and functionally distinct. This, together with the moderate heritability for phenology in Moroccan population, suggests that besides selective forces, neutral evolutionary processes have promoted intrapopulation genetic divergence and contribute to maintain high levels of genetic variation within this population. Overall, our results reinforce the importance of intrapopulation studies in long-lived plants under an evolutionary context.     

辽东栎(Quercus liaotungensis )叶特性沿海拔梯度的变化及其环境解释

植物对环境的适应一直是生态学研究的核心问题之一。山地由于海拔剧烈变化造成显著的环境差异,成为研究植物对环境适应性变化的理想对象。为阐明辽东栎（Quercus liaotungensis Koidz.）叶对环境的适应性变化,在北京东灵山地区辽东栎海拔分布范围（1000~1800m）内研究了叶特性的变化规律及其与地形和土壤养分的关系。回归分析发现：辽东栎气孔密度、气孔长度和叶面积随海拔的升高呈现曲线变化形式。在海拔约1400m处,气孔密度最小而气孔长度和叶面积最大;气孔密度和长度成反比;叶长宽比没有明显变化;叶绿素（a＋b）含量和单位干重叶氮、磷和钾含量沿海拔梯度呈上升趋势,同时叶绿素含量和叶氮含量有较弱的正相关。偏相关分析显示：叶绿素含量和坡位有显著的相关关系,叶磷含量与坡度关系显著,但叶养分与土壤养分之间未表现出明显的相关关系;地形和土壤养分与气孔密度、长度和叶面积等形态指标的关系均不显著。方差分析表明上坡位与中、下坡位的叶绿素含量有显著差异,上坡位的叶绿素含量最高。辽东栎大部分叶特性在其海拔分布范围内有显著的变化,并且形态特征和养分特征的变化形式不同,海拔1400m左右是辽东栎叶形态特征变化最显著的范围。这些叶特征的变化与土壤养分的关系不明显。     

Evolutionary dynamics of the leaf phenological cycle in an oak metapopulation along an elevation gradient

It has been predicted that environmental changes will radically alter the selective pressures on phenological traits. Long‐lived species, such as trees, will be particularly affected, as they may need to undergo major adaptive change over only one or a few generations. The traits describing the annual life cycle of trees are generally highly evolvable, but nothing is known about the strength of their genetic correlations. Tight correlations can impose strong evolutionary constraints, potentially hampering the adaptation of multivariate phenological phenotypes. In this study, we investigated the evolutionary, genetic and environmental components of the timing of leaf unfolding and senescence within an oak metapopulation along an elevation gradient. Population divergence, estimated from in situ and common‐garden data, was compared to expectations under neutral evolution, based on microsatellite markers. This approach made it possible (1) to evaluate the influence of genetic correlation on multivariate local adaptation to elevation and (2) to identify traits probably exposed to past selective pressures due to the colder climate at high elevation. The genetic correlation was positive but very weak, indicating that genetic constraints did not shape the local adaptation pattern for leaf phenology. Both spring and fall (leaf unfolding and senescence, respectively) phenology timings were involved in local adaptation, but leaf unfolding was probably the trait most exposed to climate change‐induced selection. Our data indicated that genetic variation makes a much smaller contribution to adaptation than the considerable plastic variation displayed by a tree during its lifetime. The evolutionary potential of leaf phenology is, therefore, probably not the most critical aspect for short‐term population survival in a changing climate.     

Phenotypic divergence of the common toad (Bufo bufo) along an altitudinal gradient: evidence for local adaptation

Variation in the environment can induce different patterns of genetic and phenotypic differentiation among populations. Both neutral processes and selection can influence phenotypic differentiation. Altitudinal phenotypic variation is of particular interest in disentangling the interplay between neutral processes and selection in the dynamics of local adaptation processes but remains little explored. We conducted a common garden experiment to study the phenotypic divergence in larval life-history traits among nine populations of the common toad (Bufo bufo) along an altitudinal gradient in France. We further used correlation among population pairwise estimates of quantitative trait (Q_ST) and neutral genetic divergence (F_ST from neutral microsatellite markers), as well as altitudinal difference, to estimate the relative role of divergent selection and neutral genetic processes in phenotypic divergence. We provided evidence for a neutral genetic differentiation resulting from both isolation by distance and difference in altitude. We found evidence for phenotypic divergence along the altitudinal gradient (faster development, lower growth rate and smaller metamorphic size). The correlation between pairwise Q_STs–F_STs and altitude differences suggested that this phenotypic differentiation was most likely driven by altitude-mediated selection rather than by neutral genetic processes. Moreover, we found different divergence patterns for larval traits, suggesting that different selective agents may act on these traits and/or selection on one trait may constrain the evolution on another through genetic correlation. Our study highlighted the need to design more integrative studies on the common toad to unravel the underlying processes of phenotypic divergence and its selective agents in the context of environmental clines.     

VARIATION IN LEAF DISSECTION AND LEAF ENERGY BUDGETS AMONG POPULATIONS OF ACHILLEA FROM AN ALTITUDINAL GRADIENT

The degree of leaf dissection differs dramatically among populations of the Achillea millefolium complex along an altitudinal gradient in the Sierra Nevada. The purpose of this study was to determine whether there was a genetic basis to differences in leaf shape among populations, and also to determine the importance of genetic variability within populations, plastic responses of leaf shape to the growth environment, and genetic differences among populations in plastic response to the environment. A second major goal of the research was to investigate the effects of differences in leaf dissection on the capacity for leaf temperatures to become uncoupled from air temperatures. Greenhouse experiments using clonal replicates of plants collected at different altitudes revealed that there were genetic differences among populations in the degree of dissection, and that other sources of phenotypic variation, such as plasticity, were also significant. Leaves from the highest altitude population had the most compact shape under all conditions, while those from lower altitude populations were always more open and highly dissected. In both simulations and actual measurements the dissected leaves of low altitude plants remained close to air temperatures, while the compact leaves of high altitude plants had the capacity to warm up substantially above air temperatures.     

Sources of Variation in Leaf Shape among Two Populations of Achillea Lanulosa

Achillea lanulosa has complex, highly dissected leaves that vary in shape and size along an altitudinal gradient. Plants from a high and an intermediate altitude population were clonally replicated and grown in a controlled environment at warm and cool conditions under bright light. There were genetic differences among populations and among individuals within populations in leaf size and shape. Heritabilities for leaf size and shape characters were moderate. Leaves of the lower altitude population were larger and differed from the higher altitude plants in both coarse and fine shape. Plastic response to temperature of the growth environment paralleled the genetic differentiation between low and high altitude populations. There was no apparent trade-off between genetic control over morphology and the capacity for directional plastic response to the environment. Differences in leaf dissection and size at contrasting altitudes in this species are the result of both genetic divergence among populations and of acclimative responses to local environments.     

Grasshopper populations across 2000 m of altitude: is there life history adaptation?

Life history differentiation along climatic gradients may have allowed a species to extend its geographic range. To explore this hypothesis, we compared eleven Omocestus viridulus (Orthoptera: Acrididae) populations along an altitudinal gradient from 410 to 2440 m in Switzerland, both in the field and laboratory. In situ temperature records indicated a striking decline in available heat sums along the gradient, and field populations at high altitudes reached egg hatching and adulthood much later in the year than at low elevation. The reproductive period at high altitude is thus severely limited by season length, especially during a cool year. However, controlled environment experiments revealed that intrinsic rates of embryonic and juvenile development increased with the populations' altitude of origin. This countergradient variation is largely genetic and conforms to predictions of life history theory. No corresponding differentiation in the overwintering egg stage, a pivotal determinant of phenology, was found. This trait seems conserved within the gomphocerine grasshopper subfamily. Although we found evidence for altitudinal adaptation in development, the potential of O. viridulus to adapt to cool alpine climates appears restricted by a phylogenetic constraint.     

辽东栎（Quercus liaotungensis ）叶特性沿海拔梯度的变化及其环境解释

植物对环境的适应一直是生态学研究的核心问题之一。山地由于海拔剧烈变化造成显著的环境差异,成为研究植物对环境适应性变化的理想对象。为阐明辽东栎(Quercus liaotungensis Koidz.)叶对环境的适应性变化,在北京东灵山地区辽东栎海拔分布范围(1000~1800 m)内研究了叶特性的变化规律及其与地形和土壤养分的关系。回归分析发现：辽东栎气孔密度、气孔长度和叶面积随海拔的升高呈现曲线变化形式。在海拔约1400m处,气孔密度最小而气孔长度和叶面积最大;气孔密度和长度成反比;叶长宽比没有明显变化;叶绿素(a+b)含量和单位干重叶氮、磷和钾含量沿海拔梯度呈上升趋势,同时叶绿素含量和叶氮含量有较弱的正相关。偏相关分析显示：叶绿素含量和坡位有显著的相关关系,叶磷含量与坡度关系显著,但叶养分与土壤养分之间未表现出明显的相关关系;地形和土壤养分与气孔密度、长度和叶面积等形态指标的关系均不显著。方差分析表明上坡位与中、下坡位的叶绿素含量有显著差异,上坡位的叶绿素含量最高。辽东栎大部分叶特性在其海拔分布范围内有显著的变化,并且形态特征和养分特征的变化形式不同,海拔1400 m左右是辽东栎叶形态特征变化最显著的范围。这些叶特征的变化与土壤养分的关系不明显。     

Correlated changes in thermotolerance traits and body color phenotypes in montane populations of Drosophila melanogaster: analysis of within- and between-population variations

Thermotolerance traits vary across geographical gradients but there is a lack of clinal variation in some Drosophila species. Thus, it is not clear whether thermotolerance or other correlated traits are the target of natural selection. In order to test selection responses, we investigated body melanization and thermotolerance traits in six altitudinal populations of Drosophila melanogaster . Based on rearing different geographical populations under uniform growth conditions at 21 °C (common garden experiments), clinal variations for cold resistance are in the direction opposite to heat resistance along an altitudinal gradient, that is darker flies from highland populations evidenced higher levels of cold resistance while lowland populations showed higher heat resistance. Phenotypic plastic responses for body melanization at 17–28 °C showed significant correlations with thermotolerance traits. At 17 °C, regression coefficients as a function of altitude are highly significant and positive for cold resistance but negative for heat knockdown. However, for flies reared at 28 °C, there is no elevational change in melanization as well as thermotolerance traits. Thus, both genetic and plastic changes of body melanization and thermotolerance traits suggest a correlated selection response. Further, within-population analyses of body melanization (based on dark, intermediate and light color phenotypes) showed significant associations with thermotolerance traits. Correlated variations in body melanization and thermal tolerances are associated with climatic thermal variability ( T _cv) but not with T _min. or T _max. along an altitudinal gradient.     

Population differentiation of sessile oak at the altitudinal front of migration in the French Pyrenees

To assess the effects of altitude on the level and structure of genetic diversity, a genetic survey was conducted in 12 populations of sessile oak (Quercus petraea) located between 130 and 1660 m in two parallel valleys on the northern side of the Pyrenees Mountains. Genetic diversity was monitored at 16 nuclear microsatellite loci and 5 chloroplast DNA (cpDNA) markers. The cpDNA survey suggested that extant populations in both valleys shared the same source populations from the plain. There was no visible trend of nuclear genetic diversity along altitude, even if indirect estimates of effective population sizes revealed a consistent reduction at higher altitudes. Population differentiation, although low, was mostly present among populations of the same valleys and reached similar levels than differentiation across the range of distribution of sessile oak. Contribution to the overall differentiation in the valleys was mostly due to the genetic divergence of the highest populations and the altitudinal variation of allelic frequencies at a few loci. Bayesian inference of migration between groups of populations showed that gene flow is preferentially unidirectional from lower altitudes in one valley to other groups of populations. Finally, we found evidence of clonal reproduction in high altitude populations. The introgression of Quercus robur and Quercus pubescens was also more frequent at the altitudinal margin suggesting that this mechanism may have contributed to the present migration and adaptation of Q. petraea and may also facilitate its future upslope shift in the context of climate change.     

Arabidopsis thaliana populations show clinal variation in a climatic gradient associated with altitude

? Understanding the adaptive basis of life history variation is a central goal in evolutionary ecology. The use of model species enables the combination of molecular mechanistic knowledge with ecological and evolutionary questions, but the study of life history variation in natural environments is required to merge these disciplines. ? Here, we tested for clinal variation in life history and associated traits along an environmental and altitudinal gradient in the model species Arabidopsis thaliana. Seventeen natural populations of A. thaliana were geo-referenced in north-eastern Spain on a gradient in which precipitation increases but maximum spring temperature and minimum winter temperature decrease with altitude. ? One hundred and eighty-nine genotypes from the 17 populations were grown under uniform controlled conditions. Variations in traits related to biomass allocation, fecundity, phenology and vegetative growth were tested for relationships with the altitude and climatic variables associated with the home sites. Above-ground mass, number of rosette leaves at bolting, developmental time and seed weight increased with the home site's altitude. Root allocation, vegetative growth during winter and number of seeds decreased with altitude. ? We suggest that the differences among home sites provide clues to the variation in adaptive strategies associated with the climatic gradient. We compared these results with adaptations and clinal relationships reported for other species and with molecular mechanisms described in Arabidopsis.     

Clinal differentiation during invasion: <Emphasis Type="Italic">Senecio inaequidens</Emphasis> (Asteraceae) along altitudinal gradients in Europe

The dynamics of plant population differentiation may be integral in predicting aspects of introduced species invasion. In the present study, we tested the hypothesis that European populations of Senecio inaequidens (Asteraceae), an invasive species with South African origins, differentiated during migration from two independent introduction sites into divergent altitudinal and climatic zones. We carried out 2 years of common garden experiments with eight populations sampled from Belgian and ten populations from French altitudinal transects. The Belgian transect followed a temperature and precipitation gradient. A temperature and summer drought gradient characterized the French transect. We evaluated differentiation and clinal variation in plants germinated from field-collected seed using the following traits: days to germination, days to flowering, height at maturity, final plant height and aboveground biomass. Results showed that S. inaequidens populations differentiated in growth traits during invasion. During the 1st year of sampling, the results indicated clinal variation for growth traits along both the Belgium and French altitudinal transects. Data from the 2nd year of study demonstrated that with increasing altitude, a reduction in three growth traits, including plant height at maturity, final plant height and aboveground biomass, was detected along the French transect, but no longer along the Belgian one. Phenological traits did not exhibit a clear clinal variation along altitudinal transects. The possible evolutionary causes for the observed differentiation are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Desiccation resistance along an aridity gradient in the cactophilic fly Drosophila buzzatii: sex-specific responses to stress

Stress resistance characters are valuable tools for the study of acclimation potential, adaptive strategies and biogeographic patterns in species exposed to environmental variability. Water stress is a challenge to terrestrial arthropods because of their small size and relatively high area: volume ratio. Fruit flies have been investigated to record adaptive morphological and physiological traits, as well as to test their responses to stressful factors. In this study, we investigate the ability to cope with water stress, by examining variation in desiccation resistance in a species that lives mainly in desert lands. Specifically, we explored the genetic and ecological basis of desiccation resistance in populations of Drosophila buzzatii from Northern Argentina. We used a common garden experiment with desiccation treatments on a number of isofemale lines from four populations along an aridity gradient. Our results revealed significant among-population differentiation and substantial amounts of genetic variation for desiccation resistance. We also detected significant genotype-by-environment and genotype-by-sex interactions indicative that desiccation resistance responses of the lines assayed were environment- and sex-specific. In addition, we observed clinal variation in female desiccation resistance along gradients of altitude, temperature and humidity; that desiccation resistance is a sexually dimorphic trait, and that sexual dimorphism increased along the aridity and altitudinal gradients. Based on current evidence, we propose that the observed sex-specific responses may reflect different life history traits, and survival and reproductive strategies in different ecological scenarios.