Local adaptation of stress related traits in Drosophila buzzatii and Drosophila simulans in spite of high gene flow

We addressed the question if local adaptation to a thermal gradient is possible in spite of a high gene flow among closely spaced populations of two species of Drosophila from the island of La Gomera (Canary Islands). Variation in multiple traits related to stress resistance in different life stages was measured in both species in flies collected from five localities at different altitudes and thereby with different climatic conditions. Based on microsatellite loci, the populations were not genetically differentiated. However, 18 of the 24 independent traits measured showed significant differentiation among populations of Drosophila buzzatii, but only nine of 25 for Drosophila simulans. This difference in the number of traits might reflect higher habitat specificity and thus higher potential for local adaptation of D. buzzatii than D. simulans. We found clinal variation, as some traits showed significant linear regressions on altitude, but more on altitude cubed.     

Altitudinal variation for stress resistance traits and thermal adaptation in adult Drosophila buzzatii from the New World

Multiple stress resistance traits were investigated in the cactophilic fly Drosophila buzzatii. Adults from seven populations derived from North-Western Argentina were compared with respect to traits relevant for thermal stress resistance and for resistance to other forms of environmental stress. The populations were collected along an altitudinal gradient spanning more than 2000 m in height, showing large climatic differences. The results suggest that knock-down resistance to heat stress, desiccation resistance and Hsp70 expression at a relatively severe stressful temperature best reflect thermal adaptation in this species. Furthermore, cold resistance seemed to be of less importance than heat resistance, at least for the adult life stage, in these populations. Clinal variation in thermal resistance traits over short geographical distances suggests relatively strong adaptive differentiation of the populations. This study provides the first evidence for altitudinal differentiation in stress-related traits, and suggests that Hsp70 expression level can be related to altitudinal clines of heat-stress resistance.     

Rapid adaptive evolution of photoperiodic response during invasion and range expansion across a climatic gradient

Abstract Understanding the mechanisms of adaptation to spatiotemporal environmental variation is a fundamental goal of evolutionary biology. This issue also has important implications for anticipating biological responses to contemporary climate warming and determining the processes by which invasive species are able to spread rapidly across broad geographic ranges. Here, we compare data from a historical study of latitudinal variation in photoperiodic response among Japanese and U.S. populations of the invasive Asian tiger mosquito Aedes albopictus with contemporary data obtained using comparable methods. Our results demonstrated rapid adaptive evolution of the photoperiodic response during invasion and range expansion across ～15° of latitude in the United States. In contrast to the photoperiodic response, size-based morphological traits implicated in climatic adaptation in a wide range of other insects did not show evidence of adaptive variation in Ae. albopictus across either the U.S. (invasive) or Japanese (native) range. These results show that photoperiodism has been an important adaptation to climatic variation across the U.S. range of Ae. albopictus and, in conjunction with previous studies, strongly implicate the photoperiodic control of seasonal development as a critical evolutionary response to ongoing contemporary climate change. These results also emphasize that photoperiodism warrants increased attention in studies of the evolution of invasive species.     

Reproductive diapause and life-history clines in North American populations of Drosophila melanogaster

Latitudinal clines are widespread in Drosophila melanogaster, and many have been interpreted as adaptive responses to climatic variation. However, the selective mechanisms generating many such patterns remain unresolved, and there is relatively little information regarding how basic life-history components such as fecundity, life span and mortality rates vary across environmental gradients. Here, it is shown that four life-history traits vary predictably with geographic origin of populations sampled along the latitudinal gradient in the eastern United States. Although such patterns are indicative of selection, they cannot distinguish between the direct action of selection on the traits in question or indirect selection by means of underlying genetic correlations. When independent suites of traits covary with geography, it is therefore critical to separate the widespread effects of population source from variation specifically for the traits under investigation. One trait that is associated with variation in life histories and also varies with latitude is the propensity to express reproductive diapause; diapause expression has been hypothesized as a mechanism by which D. melanogaster adults overwinter, and as such may be subject to strong selection in temperate habitats. In this study, recently derived isofemale lines were used to assess the relative contributions of population source and diapause genotype in generating the observed variance for life histories. It is shown that although life span, fecundity and mortality rates varied predictably with geography, diapause genotype explained the majority of the variance for these traits in the sampled populations. Both heat and cold shock resistance were also observed to vary predictably with latitude for the sampled populations. Cold shock tolerance varied between diapause genotypes and the magnitude of this difference varied with geography, whereas heat shock tolerance was affected solely by geographic origin of the populations. These data suggest that a subset of life-history parameters is significantly influenced by the genetic variance for diapause expression in natural populations, and that the observed variance for longevity and fecundity profiles may reflect indirect action of selection on diapause and other correlated traits.     

Geographical Clines for Quantitative Traits in Natural Populations of a Tropical Drosophilid: Zaprionus Indianus

We analyzed natural populations of Zaprionus indianusin 10 Indian localities along a south-north transect (latitude: 10–31°3 N). Size traits (body weight, wing length and thorax length) as well as a reproductive trait (ovariole number) followed a pattern of clinal variation, that is, trait value increased with latitude. Wing/thorax ratio, which is inversely related to wing loading, also had a positive, but non-significant correlation with latitude. By contrast, bristle numbers (sternopleural and abdominal) exhibited a non-significant but negative correlation with latitude. Sex dimorphism, estimated as the female/male ratio, was very low in Z. indianus, contrasting with results already published in other species. Genetic variations among populations were also analyzed according to other geographic parameters (altitude and longitude) and to climatic conditions from each locality. A significant effect of altitude was found for size traits. For abdominal bristles, a multiple regression technique evidenced a significant effect of both latitude and altitude, but in opposite directions. Genetic variations were also correlated to climate, and mainly with average year temperature. Taking seasonal variations into account failed however to improve the predictability of morphometrical variations. The geographic differentiation of Z.indianusfor quantitative traits suggests adaptive response to local conditions, especially to temperature, but also reveals a complex situation according to traits investigated and to environmental parameters, which does not match results on other drosophilid species.     

Root architecture,plant size and soil nutrient variation in natural populations of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Phenotypic variation in ecologically important traits may vary at large and small geographic scales, and may be shaped by natural selection. Here our explicit aim is to evaluate phenotypic differentiation among local populations and examine its relationship with ecological edaphic and climatic features that could lead to local adaptation. We characterized six populations of the model plant Arabidopsis thaliana over 3 years in the field in its native range. At each site, we measured edaphic conditions and aboveground and belowground phenotypes. In addition, we grew plants from the six characterized populations in a common greenhouse along with an additional fifteen populations from the Iberian Peninsula to examine evolutionary and ecological differentiation among populations, and relationships between geographic and ecological distance to phenotypic differences among populations. Significant differences in aboveground and belowground traits, population density, and micro- and macro-nutrient soil concentrations were found among the field populations. In particular, root architectural traits differed significantly among field populations. Complex patterns of ecological differences among population and plant phenotypes emerged when examining edaphic conditions in the Extremadura region, and geographic and climate variables at a broader scale of the Iberian Peninsula. We report levels of phenotypic variation at the local scale comparable to those found at broad geographic scales and report that local edaphic conditions contribute to population-level phenotypic variation in root and shoot traits. To our knowledge, these are the first reports of among population root architectural variation from natural field populations for this model organism. We demonstrate how ecological features, such as soil nutrients, can be associated with the phenotypic variation of A. thaliana measured in natural populations and may contribute to adaptive differentiation at a local scale.     

Patterns of Cross-Continental Variation in Tree Seed Mass in the Canadian Boreal Forest

Seed mass is an adaptive trait affecting species distribution, population dynamics and community structure. In widely distributed species, variation in seed mass may reflect both genetic adaptation to local environments and adaptive phenotypic plasticity. Acknowledging the difficulty in separating these two aspects, we examined the causal relationships determining seed mass variation to better understand adaptability and/or plasticity of selected tree species to spatial/climatic variation. A total of 504, 481 and 454 seed collections of black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss) and jack pine (Pinus banksiana Lamb) across the Canadian Boreal Forest, respectively, were selected. Correlation analyses were used to determine how seed mass vary with latitude, longitude, and altitude. Structural Equation Modeling was used to examine how geographic and climatic variables influence seed mass. Climatic factors explained a large portion of the variation in seed mass (34, 14 and 29%, for black spruce, white spruce and jack pine, respectively), indicating species-specific adaptation to long term climate conditions. Higher annual mean temperature and winter precipitation caused greater seed mass in black spruce, but annual precipitation was the controlling factor for white spruce. The combination of factors such as growing season temperature and evapotranspiration, temperature seasonality and annual precipitation together determined seed mass of jack pine. Overall, sites with higher winter temperatures were correlated with larger seeds. Thus, long-term climatic conditions, at least in part, determined spatial variation in seed mass. Black spruce and Jack pine, species with relatively more specific habitat requirements and less plasticity, had more variation in seed mass explained by climate than did the more plastic species white spruce. As traits such as seed mass are related to seedling growth and survival, they potentially influence forest species composition in a changing climate and should be included in future modeling of vegetation shifts.     

Polymorphic inversion and esterase loci complex on chromosome 2 of Drosophila buzzatii. II. Spatial variation

The potential influence of linked inversions on allele frequency variation at the Est-1 and Est-2 loci among Australian populations of D. buzzatii was determined by statistical analyses allele and inversion gametic frequencies. Most of the significant spatial and climatic associations found for all Est-1 allele frequencies, and for one allele only of Est-2 (Est-2c+), were accounted for by their linkage disequilibria with the inversions, which covaried with environmental variables. Consistent with this result, the spatial and climatic associations for conditional Est-1 and Est-2 allele frequencies tended to be different from those for the respective unadjusted allele frequencies. In one important respect, the results for Est-1 and Est-2 were not altered by inversions. For both unadjusted and conditional Est-1 allele frequencies, few climatic associations remain after correcting for geographic location, whereas for both unadjusted and conditional Est-2 allele frequencies, climatic associations remain after correcting for geographic location. Thus, apparent selection affecting allele frequencies at the Est-2 locus is not accounted for by linked inversions.     

An adaptive explanation for geographically structured allozyme variation inDichroplus elongatus (Orthoptera: Acrididae)

Previous studies on allozyme variation in five populations of the grasshopperDichroplus elongatus along a geographical gradient in Argentina revealed a significant degree of population structuring and a significant association between one of the loci (Aat-1) and latitude. As this could not be entirely explained by historical factors, the possible adaptive significance of this locus or loci in linkage disequilibrium was investigated in the present study, with an emphasis on the role of environmental variables correlated with latitude. The present paper reports a study of the geographical organization of allozyme diversity over a wider range of the distribution ofD. elongatus. The relation of allelic frequencies with geographic climatic variables was analysed. We have found (i) that different loci (Aat-1 andPep-1) covary significantly with different variables, (ii) discordance between genetic and geographical distances, (iii) copious gene flow that would mask allelic frequency differences due solely to genetic drift, (iv) and temporal stability of the gradient found. The results suggest that causes other than drift and migration may explain the observed directional patterns of variation inD. elongatus.     

Genetic and morphological differentiation in Populus nigra L.: isolation by colonization or isolation by adaptation?

Identifying processes underlying the genetic and morphological differences among populations is a central question of evolutionary biology. Forest trees typically contain high levels of neutral genetic variation, and genetic differences are often correlated with geographic distance between populations [isolation by distance (IBD)] or are due to historic vicariance events [isolation by colonization (IBC)]. In contrast, morphological differences are largely due to local adaptation. Here, we examined genetic (microsatellite) and morphological (from a common garden experiment) variation in Populus nigra L., European black poplar, collected from 13 sites across western Europe and grown in a common garden in Belgium. Significant genetic differentiation was observed, with populations from France displaying greater admixture than the distinct Spanish and central European gene pools, consistent with previously described glacial refugia (IBC). Many quantitative traits displayed a bimodal distribution, approximately corresponding to small‐leaf and large‐leaf ecotypes. Examination of nine climatic variables revealed the sampling locations to have diverse climates, and although the correlation between morphological and climatic differences was significant, the pattern was not consistent with strict local adaptation. Partial Mantel tests based on multivariate summary statistics identified significant residual correlation in comparisons of small‐leaf to large‐leaf ecotypes, and within the small‐leaf samples, but not within large‐leaf ecotypes, indicating that variation within the small‐leaf morphotype in particular may be adaptive. Some small‐leaf populations experience climates very similar to those in large‐leaf sites. We conclude that adaptive differentiation and persistent IBC acted in combination to produce the genetic and morphological patterns observed in P. nigra.     

The ecology of an adaptive radiation of three‐spined stickleback from North Uist,Scotland

There has been a large focus on the genetics of traits involved in adaptation, but knowledge of the environmental variables leading to adaptive changes is surprisingly poor. Combined use of environmental data with morphological and genomic data should allow us to understand the extent to which patterns of phenotypic and genetic diversity within a species can be explained by the structure of the environment. Here, we analyse the variation of populations of three‐spined stickleback from 27 freshwater lakes on North Uist, Scotland, that vary greatly in their environment, to understand how environmental and genetic constraints contribute to phenotypic divergence. We collected 35 individuals per population and 30 abiotic and biotic environmental parameters to characterize variation across lakes and analyse phenotype–environment associations. Additionally, we used RAD sequencing to estimate the genetic relationships among a subset of these populations. We found a large amount of phenotypic variation among populations, most prominently in armour and spine traits. Despite large variation in the abiotic environment, namely in ion composition, depth and dissolved organic Carbon, more phenotypic variation was explained by the biotic variables (presence of predators and density of predator and competitors), than by associated abiotic variables. Genetic structure among populations was partly geographic, with closer populations being more similar. Altogether, our results suggest that differences in body shape among stickleback populations are the result of both canalized genetic and plastic responses to environmental factors, which shape fish morphology in a predictable direction regardless of their genetic starting point.     

Relationships between abiotic variables and geographic variation in skulls of pumas (Puma concolor: Mammalia, Felidae) in North and South America

Relationships between patterns of geographic variation in cranial morphology and selected abiotic variables were studied in the puma ( Puma concolor Linnaeus). Our dataset consisted of 11 cranial, 6 dental, and 2 mandibular measurements of 1700 adults, which were separated by gender and age class and analysed by univariate and multivariate statistical procedures. Variation in size was correlated with precipitation, but size was more highly correlated with latitude and temperature. The pattern of geographic variation in size of P. concolor is consistent with Bergmann's rule; populations with larger pumas occur more distant from the equator than populations with smaller pumas. A combination of climatic and biotic factors contribute to patterns of geographic variation in size of P. concolor in North and South America.     

Climatic adaptation in an isolated and genetically impoverished amphibian population

The capacity of populations to respond adaptively to environmental change is essential for their persistence. Isolated populations often harbour reduced genetic variation, which is predicted to decrease adaptive potential, and can be detrimental under the current scenarios of global change. In this study, we examined climatic adaptation in larval life history traits in the pool frog Rana lessonae along a latitudinal gradient across the northern distribution area of the species, paying special attention to the isolated and genetically impoverished fringe populations in central Sweden. Larvae from eight populations within three geographic areas (Poland, Latvia and Sweden) were reared under three temperatures (19, 22 and 26°C) in a common garden laboratory experiment. We found clear evidence for latitudinal adaptation in R. lessonae populations, larvae from higher latitudes growing and developing faster than low‐latitude ones. Larvae from the Swedish populations were able to compensate for the effects of cooler temperatures and a shorter growth season with genetically higher growth and development rates (i.e. countergradient variation) in the two higher temperature treatments, but there was no difference among the populations at the lowest temperature treatment, which is likely to be close to the temperature limiting growth in R. lessonae. Our results demonstrate that isolated and genetically impoverished populations can be locally adapted, and identify the Swedish fringe populations as a significant conservation unit adapted to the northern environmental conditions.     

Plasticity in latitudinal patterns of leaf N and P of Oryza rufipogon in China

Characterising the adaptability in nature of plant stoichiometric patterns across geographic or environmental gradients is important in advancing our understanding of the organisation of plant–nutrient relationships. We examined correlations between plant nutrient traits, latitude, longitude, climate and soil variables in 34 populations of Oryza rufipogon across its range. We further compared the responses of population transplants at two experimental gardens: one beyond its northern natural range and another near the southern limit, to assess the nature of geographic variation in plant nutrients. The study showed that leaf P of O. rufipogon in the field was negatively correlated with latitude and largely depended on temperature and soil P availability. Leaf N was not related to latitude but was significantly correlated with precipitation and soil N concentration. Leaf N:P ratio was largely determined by absorption efficiency of P. Transplantation revealed that there were no significant associations of leaf nutrients with geographic, climatic or soil variables of origin in either of the experimental gardens, indicating phenotypic plasticity. However, examination of relationships between response ratios of leaf nutrients and change ratio of climate and soil environments, as well as norms of reaction in the transplantation experiment, revealed more complexity, suggesting both substantial genotypic diversity and the existence of genotype × environment interactions in these populations of O. rufipogon. These data indicate that adaptive plasticity response of plants to temperature and soil P availability significantly explain the observed shifts in leaf N, P and N:P of O. rufipogon along latitudinal gradients.     

Effect of climatic variation on the morphological characteristics of 37‐year‐old balsam fir provenances planted in a common garden in New Brunswick,Canada

The extent of the effect of projected changes in climate on trees remains unclear. This study investigated the effect of climatic variation on morphological traits of balsam fir [Abies balsamea (L.) Mill.] provenances sourced from locations spanning latitudes from 44° to 51°N and longitudes from 53° to 102°W across North America, growing in a common garden in eastern Canada. Lower latitude provenances performed significantly better than higher latitude provenances (p < .05) with regard to diameter at breast height (DBH), height (H), and crown width (CW), a distinction indicative of genotypic control of these traits. There was, however, no significant difference among provenances in terms of survival (p > .05), an indication of a resource allocation strategy directed at survival relative to productivity in higher latitude provenances as seen in their lower DBH, H, and CW compared to the lower latitude provenances. Temperature had a stronger relationship with DBH, H, and CW than precipitation, a reflection of adaptation to local conditions in populations of the species along latitudinal gradients. Both climatic variables had some effect on tree survival. These results suggest that the response of balsam fir to climatic variation will likely not be uniform in the species, but differ based on genetic characteristics between populations located in the northern and southern parts of the species’ range. Population differences in response to climatic variation may be evident earlier in growth traits, compared to survival in balsam fir. The findings of this study will facilitate modeling in the species that is reflective of genetic variation in response to climatic conditions, and guide provenance selection for utilization in terms of productivity or resilience as well as breeding programs directed at obtaining species that possibly combine both traits.     

Latitudinal patterns in rodent metabolic flexibility

Macrophysiology is defined as the study of variation in physiological traits-including physiological trait flexibility-over large geographical and temporal scales, and the ecological implications of this variation. A classic example of a macrophysiological trend is the one emerging from the climatic variability hypothesis, which states that as the range of climatic fluctuation experienced by terrestrial animals increases with latitude, individuals at higher latitudes should be more plastic than individuals inhabiting lower latitudes. In this context, we evaluate the correlation between absolute metabolic scope during cold exposure (an instantaneous measure of metabolic flexibility) and different geographic and climatic variables for 48 rodent species. Conventional and phylogenetic informed analyses indicated a positive correlation between metabolic scope and geographic latitude. These findings, together with previous reports on latitudinal pattern in phenotypic flexibility, suggest that an increase in physiological flexibility with latitude may hold for many phenotypic traits.     

Covariation of larval gene expression and adult body size in natural populations of Drosophila melanogaster

Understanding adaptive phenotypic variation is one of the most fundamental problems in evolutionary biology. Genes involved in adaptation are most likely those that affect traits most intimately connected to fitness: life-history traits. The genetics of quantitative trait variation (including life histories) is still poorly understood, but several studies suggest that (1) quantitative variation might be the result of variation in gene expression, rather than protein evolution, and (2) natural variation in gene expression underlies adaptation. The next step in studying the genetics of adaptive phenotypic variation is therefore an analysis of naturally occuring covariation of global gene expression and a life-history trait. Here, we report a microarray study addressing the covariation in larval gene expression and adult body weight, a life-history trait involved in adaptation. Natural populations of Drosophila melanogaster show adaptive geographic variation in adult body size, with larger animals at higher latitudes. Conditions during larval development also affect adult size with larger flies emerging at lower temperatures. We found statistically significant differences in normalized larval gene expression between geographic populations at one temperature (genetic variation) and within geographic populations between temperatures (developmental plasticity). Moreover, larval gene expression correlated highly with adult weight, explaining 81% of its natural variation. Of the genes that show a correlation of gene expression with adult weight, most are involved in cell growth or cell maintenance or are associated with growth pathways.     

Variation in eggshell traits between geographically distant populations of pied flycatchers Ficedula hypoleuca

The expression and impact of maternal effects may vary greatly between populations and environments. However, little is known about large‐scale geographical patterns of variation in maternal deposition to eggs. In birds, as in other oviparous animals, the outermost maternal component of an egg is the shell, which protects the embryo, provides essential mineral resources and allows its interaction with the environment in the form of gas exchange. In this study, we explored variation of eggshell traits (mass, thickness, pore density and pigmentation) across 15 pied flycatcher populations at a large geographic scale. We found significant between‐population variation in all eggshell traits, except in pore density, suggesting spatial variation in their adaptive benefits or in the females’ physiological limitations during egg laying. Between‐ population variation in shell structure was not due to geographic location (latitude and longitude) or habitat type. However, eggshells were thicker in populations that experienced higher ambient temperature during egg laying. This could be a result of maternal resource allocation to the shell being constrained under low temperatures or of an adaptation to reduce egg water loss under high temperatures. We also found that eggshell colour intensity was positively associated with biliverdin pigment concentration, shell thickness and pore density. To conclude, our findings reveal large‐ scale between‐population variation of eggshell traits, although we found little environmental dependency in their expression. Our findings call for further studies that explore other environmental factors (e.g. calcium availability and pollution levels) and social factors like sexual selection intensity that may account for differences in shell structure between populations.     

Genetically based latitudinal variation in Artemisia californica secondary chemistry

Steep climatic gradients may select for clinal adaptation in plant functional traits with implications for interspecific interactions and response to future climate change. Terpenes are common in Mediterranean environments and mediate plant interactions with both the abiotic and biotic environment, including herbivores. Clines in traits such as terpenes have received much attention because they are linked to plant fitness and experience strong selection from the abiotic and biotic environment. In this study, we tested for intraspecific variation in Artemisia californica terpene chemistry in a common garden of plants sourced from populations spanning a large precipitation gradient (6° latitude) and grown in treatments of high and low precipitation. We found genetic variation in terpene richness, diversity, concentration and composition among A. californica populations spanning this species’ range. Of these traits, terpene composition and monoterpene concentration varied clinally with respect to source site latitude. Regarding terpene composition, pairwise dissimilarity among populations increased in parallel with geographic distance between source sites. At the same time, monoterpene concentration decreased monotonically from plants of southern origin (source sites with high temperature, aridity, and precipitation variability) to plants of northern origin. Our precipitation manipulation suggests that phenotypic selection by precipitation may underlie this clinal variation in monoterpene concentration, and that monoterpene concentration and other aspects of terpene chemistry are not phenotypically plastic. In summary, this study provides novel evidence for a genetically based latitudinal cline in plant secondary chemistry and suggests that adaptation to a key aspect of the abiotic environment may contribute to this intraspecific variation. Accordingly, changes in terpene chemistry under projected future climates will likely occur solely through the relatively slow process of adaptation, with important consequences for plant interactions with the abiotic environment and a diverse community of associates.     

Postponed reproduction as an adaptation to winter conditions in Drosophila melanogaster: evidence for clinal variation under semi-natural conditions

Patterns of climatic adaptation in drosophila and other insects have largely been inferred from laboratory comparisons of traits that vary clinally. Here, we extend this research to comparisons under semi-natural conditions. To test for clinal variation in reproductive patterns and survival over winter, Drosophila melanogaster populations were initiated from seven collection sites along the eastern coast of Australia, ranging from tropical to temperate regions. The fecundity and survival of these populations were monitored in field cages at a temperate location until all adults had died more than 5 months later. Total fecundity showed a curvilinear relationship with latitude, due to higher egg production by high- and low-latitude populations. Adults from temperate locations survived winter conditions better than those from subtropical populations but not tropical ones. There was a linear cline in the timing of egg production: temperate populations produced eggs later than populations from lower latitudes. This cline is likely to be adaptive because egg-to-adult viability experiments indicated that only eggs laid in spring developed successfully to the adult stage. There was no evidence for climatic adaptation in the immature stages. The adult mortality rate increased gradually over winter, and in some populations was also correlated with the minimum ambient temperature. These results indicate that adaptation to winter conditions in D. melanogaster has involved shifts in reproductive patterns.