Environmental Clines Research Articles

Allelic variants confer Arabidopsis adaptation to small regional environmental differences.

Natural populations of Arabidopsis thaliana provide powerful systems to study the adaptation of wild plant species. Previous research has predominantly focused on global populations or accessions collected from regions with diverse climates. However, little is known about the genetics underlying adaptation in regions with mild environmental clines. We have examined a diversity panel consisting of 192 A. thaliana accessions collected from the Netherlands, a region with limited climatic variation. Despite the relatively uniform climate, we identified evidence of local adaptation within this population. Notably, semidwarf accessions, due to mutation of the GIBBERELLIC ACID REQUIRING 5 (GA5) gene, occur at a relatively high frequency near the coast and these displayed enhanced tolerance to high wind velocities. Additionally, we evaluated the performance of the population under iron deficiency conditions and found that allelic variation in the FE SUPEROXIDE DISMUTASE 3 (FSD3) gene affects tolerance to low iron levels. Moreover, we explored patterns of local adaptation to environmental clines in temperature and precipitation, observing that allelic variation at LA RELATED PROTEIN 1C (LARP1c) likely affects drought tolerance. Not only is the genetic variation observed in a diversity panel of A. thaliana collected in a region with mild environmental clines comparable to that in collections sampled over larger geographic ranges but it is also sufficiently rich to elucidate the genetic and environmental factors underlying natural plant adaptation.

Genetic monitoring uncovers long-distance marine feeding coupled with strong spatial segregation in sea trout (Salmo trutta L.) consistent at annual and decadal time scales

Abstract Genetic data have greatly increased means to understand fish marine migration behaviours at large spatial scale within a quantitative framework. The anadromous sea trout is a prized target of recreational fishery and an important ecosystem component in freshwater and marine coastal habitats in large parts of temperate northern Europe. Nonetheless, little is known about population distributions while feeding at sea. To reconcile notions about feeding migrations being predominantly locally restricted or not, we used SNP data for 3465 trout representing >100 rivers to identify population origins of 903 coastally feeding fish captured throughout the North Sea-Baltic Sea transition area. Across areas, coastal collections generally showed marked stock-mixing and overall 70:30% native:non-native fish with seasonal variation corresponding with spawning run timing. Data revealed strong spatial feeding segregation between trout from the Scandinavian Peninsula and the European continent. This is surprising given the short distances between areas, but is in alignment with strong genetic differentiation between populations in these areas. Estimation of stock complexity of coastal feeding aggregations showed no spatial trend through the North Sea-Baltic Sea transition area otherwise characterized by strong environmental clines. Analyses of scale samples collected in the 1950s indicate that stock-mixing was consistent over almost 70 years.

Genetic and phenotypic diversification in a widespread fish, the Sailfin Molly (Poecilia latipinna)

Widespread species often experience significant environmental clines over the area they naturally occupy. We investigated a widespread livebearing fish, the Sailfin molly (Poecilia latipinna) combining genetic, life-history, and environmental data, asking how structured populations are. Sailfin mollies can be found in coastal freshwater and brackish habitats from roughly Tampico, Veracruz in Mexico to Wilmington, North Carolina, in the USA. In addition, they are found inland on the Florida peninsula. Using microsatellite DNA, we genotyped 168 individuals from 18 populations covering most of the natural range of the Sailfin molly. We further determined standard life-history parameters for both males and females for these populations. Finally, we measured biotic and abiotic parameters in the field. We found six distinct genetic clusters based on microsatellite data, with very strong indication of isolation by distance. However, we also found significant numbers of migrants between adjacent populations. Despite genetic structuring we did not find evidence of cryptic speciation. The genetic clusters and the migration patterns do not match paleodrainages. Life histories vary between populations but not in a way that is easy to interpret. We suggest a role of humans in migration in the sailfin molly, for example in the form of a ship channel that connects southern Texas with Louisiana which might be a conduit for fish migration.

Spatio‐temporal shift in body size and plumage coloration is associated with the magnitude of climate change in a cosmopolitan owl

AbstractAimClimatic gradients shape geographic variation in phenotypic traits that are involved in animal thermoregulation. Therefore, increasing temperatures under global warming are expected to cause change over time in traits that show predictable spatial patterns according to environmental clines (body and appendage size, tegument coloration), as well exemplified by biogeographic rules and shown by increasing literature at a local scale. However, whether temporal shifts in phenotypic traits vary spatially according to the magnitude of the change in local climatic conditions is still unknown.LocationWorld.TaxonBarn owl species complex (Tyto alba group).MethodsBy using thousands of museum specimens collected across the globe from 1901 to 2018, we calculated within‐population change over time in wing length, bill length and melanin‐based plumage coloration and examined whether these trends differed across geographic regions, latitudes and gradients of climate changes.ResultsWe showed that populations exposed to an increasing warmer climate were subjected to a more marked decrease in body size, as gauged by wing length, and also absolute bill length that displayed a similar, although less steep, pattern of shrinking over time. In addition, phaeomelanin‐based ventral plumage colour has become paler in regions where the climate changed into warmer and drier, and darker where both temperature and precipitation increased.Main ConclusionsThese trends are generally coherent with the predictions based on Bergmann's and Gloger's rules, but not Allen's rule, and suggest that temporal shifts in body size and tegument colour depend on the magnitude of the alteration in climatic conditions, with populations living in regions where the climate has changed the most that are subjected to more marked phenotypic changes. Spatio‐temporal variation in climate has driven within‐species phenotypic clines, and it is expected to increase differences among populations according to the predicted further climate change.

Genomic responses to climate: Understanding local adaptation in the Andean tree species Nothofagus pumilio and implications for a changing world

Societal Impact StatementForest trees tend to be strongly genetically adapted to their local environments, but climate change will probably subject trees to novel combinations of precipitation, temperature, and photoperiod. Local adaptation was investigated in the ecologically and economically important Patagonian tree species Nothofagus pumilio by characterizing its genetic diversity in relation to the varied environmental conditions across its range. These insights are useful for conservation and management decisions, for example by identifying suitable populations to establish seed source plantations for restoration and characterizing relationships with environmental drivers of selection to better understand how this species will respond to climate change.SummaryNothofagus pumilio is a foundation tree species that inhabits a 2000‐km‐long range in the southern Andes, a region with two perpendicular environmental gradients: temperature and photoperiod (North–South), and precipitation (West–East). We investigated local adaptation patterns by searching for relationships between environmental clines and signatures of adaptation in candidate genes related to stress response, growth, and phenology. Using a paired site sampling design within a landscape genome analysis, we analyzed 493 adult N. pumilio trees in 20 sampling sites across the species' latitudinal range. We screened 47,336 single nucleotide polymorphism (SNP) loci in 1632 contigs (i.e., coding regions along the genome). Population structure and genetic diversity analyses preceded four genome scan analyses using genetic and environmental data. Population structure and genetic diversity are mainly oriented along the latitude axis. Genome scans identified 445 outlier SNPs, which are loci showing signatures of selection. Temperature and photoperiod variables were associated with notably more outliers than precipitation. However, the most frequent biological functions among genes were water deprivation response and cold response, suggesting that stress response is comprised of complex and polygenic traits that are affected by many environmental variables. Our findings suggest that N. pumilio shows signatures of local adaptation to extant climate conditions, including temperature, photoperiod, and precipitation. However, climate change is likely to alter existing relationships among environmental conditions to which this species is currently adapted. These changes may have unpredictable consequences for the species' future survival, adaptation potential, and the people who depend upon these forests.

Saccharomycotina yeasts defy long-standing macroecological patterns

The Saccharomycotina yeasts ("yeasts" hereafter) are a fungal clade of scientific, economic, and medical significance. Yeasts are highly ecologically diverse, found across a broad range of environments in every biome and continent on earth; however, little is known about what rules govern the macroecology of yeast species and their range limits in the wild. Here, we trained machine learning models on 12,816 terrestrial occurrence records and 96 environmental variables to infer global distribution maps at ~1 km2 resolution for 186 yeast species (~15% of described species from 75% of orders) and to test environmental drivers of yeast biogeography and macroecology. We found that predicted yeast diversity hotspots occur in mixed montane forests in temperate climates. Diversity in vegetation type and topography were some of the greatest predictors of yeast species richness, suggesting that microhabitats and environmental clines are key to yeast diversity. We further found that range limits in yeasts are significantly influenced by carbon niche breadth and range overlap with other yeast species, with carbon specialists and species in high-diversity environments exhibiting reduced geographic ranges. Finally, yeasts contravene many long-standing macroecological principles, including the latitudinal diversity gradient, temperature-dependent species richness, and a positive relationship between latitude and range size (Rapoport's rule). These results unveil how the environment governs the global diversity and distribution of species in the yeast subphylum. These high-resolution models of yeast species distributions will facilitate the prediction of economically relevant and emerging pathogenic species under current and future climate scenarios.

Temperature and nutrition do not interact to shape the evolution of metabolic rate.

Metabolic cold adaptation, or Krogh's rule, is the controversial hypothesis that predicts a monotonically negative relationship between metabolic rate and environmental temperature for ectotherms living along thermal clines measured at a common temperature. Macrophysiological patterns consistent with Krogh's rule are not always evident in nature, and experimentally evolved responses to temperature have failed to replicate such patterns. Hence, temperature may not be the sole driver of observed variation in metabolic rate. We tested the hypothesis that temperature, as a driver of energy demand, interacts with nutrition, a driver of energy supply, to shape the evolution of metabolic rate to produce a pattern resembling Krogh's rule. To do this, we evolved replicate lines of Drosophila melanogaster at 18, 25 or 28°C on control, low-calorie or low-protein diets. Contrary to our prediction, we observed no effect of nutrition, alone or interacting with temperature, on adult female and male metabolic rates. Moreover, support for Krogh's rule was only in females at lower temperatures. We, therefore, hypothesize that observed variation in metabolic rate along environmental clines arises from the metabolic consequences of environment-specific life-history optimization, rather than because of the direct effect of temperature on metabolic rate. This article is part of the theme issue 'The evolutionary significance of variation in metabolic rates'.

Multiple genomic solutions for local adaptation in two closely related species (sheep and goats) facing the same climatic constraints.

The question of how local adaptation takes place remains a fundamental question in evolutionary biology. The variation of allele frequencies in genes under selection over environmental gradients remains mainly theoretical and its empirical assessment would help understanding how adaptation happens over environmental clines. To bring new insights to this issue we set up a broad framework which aimed to compare the adaptive trajectories over environmental clines in two domesticated mammal species co-distributed in diversified landscapes. We sequenced the genomes of 160 sheep and 161 goats extensively managed along environmental gradients, including temperature, rainfall, seasonality and altitude, to identify genes and biological processes shaping local adaptation. Allele frequencies at putatively adaptive loci were rarely found to vary gradually along environmental gradients, but rather displayed a discontinuous shift at the extremities of environmental clines. Of the 430 candidate adaptive genes identified, only 6 were orthologous between sheep and goats and those responded differently to environmental pressures, suggesting different putative mechanisms involved in local adaptation in these two closely related species. Interestingly, the genomes of the 2 species were impacted differently by the environment, genes related to signatures of selection were most related to altitude, slope and rainfall seasonality for sheep, and summer temperature and spring rainfall for goats. The diversity of candidate adaptive pathways may result from a high number of biological functions involved in the adaptations to multiple eco-climatic gradients, and a differential role of climatic drivers on the two species, despite their co-distribution along the same environmental gradients. This study describes empirical examples of clinal variation in putatively adaptive alleles with different patterns in allele frequency distributions over continuous environmental gradients, thus showing the diversity of genetic responses in adaptive landscapes and opening new horizons for understanding genomics of adaptation in mammalian species and beyond.

Quantifying the role of genome size and repeat content in adaptive variation and the architecture of flowering time in Amaranthus tuberculatus.

Genome size variation, largely driven by repeat content, is poorly understood within and among populations, limiting our understanding of its significance for adaptation. Here we characterize intraspecific variation in genome size and repeat content across 186 individuals of Amaranthus tuberculatus, a ubiquitous native weed that shows flowering time adaptation to climate across its range and in response to agriculture. Sequence-based genome size estimates vary by up to 20% across individuals, consistent with the considerable variability in the abundance of transposable elements, unknown repeats, and rDNAs across individuals. The additive effect of this variation has important phenotypic consequences-individuals with more repeats, and thus larger genomes, show slower flowering times and growth rates. However, compared to newly-characterized gene copy number and polygenic nucleotide changes underlying variation in flowering time, we show that genome size is a marginal contributor. Differences in flowering time are reflected by genome size variation across sexes and marginally, habitats, while polygenic variation and a gene copy number variant within the ATP synthesis pathway show consistently stronger environmental clines than genome size. Repeat content nonetheless shows non-neutral distributions across the genome, and across latitudinal and environmental gradients, demonstrating the numerous governing processes that in turn influence quantitative genetic variation for phenotypes key to plant adaptation.

Local adaptation with gene flow in a highly dispersive shark.

Adaptive divergence in response to environmental clines are expected to be common in species occupying heterogeneous environments. Despite numerous advances in techniques appropriate for non-model species, gene-environment association studies in elasmobranchs are still scarce. The bronze whaler or copper shark (Carcharhinus brachyurus) is a large coastal shark with a wide distribution and one of the most exploited elasmobranchs in southern Africa. Here, we assessed the distribution of neutral and adaptive genomic diversity in C. brachyurus across a highly heterogeneous environment in southern Africa based on genome-wide SNPs obtained through a restriction site-associated DNA method (3RAD). A combination of differentiation-based genome-scan (outflank) and genotype-environment analyses (redundancy analysis, latent factor mixed models) identified a total of 234 differentiation-based outlier and candidate SNPs associated with bioclimatic variables. Analysis of 26,299 putatively neutral SNPs revealed moderate and evenly distributed levels of genomic diversity across sites from the east coast of South Africa to Angola. Multivariate and clustering analyses demonstrated a high degree of gene flow with no significant population structuring among or within ocean basins. In contrast, the putatively adaptive SNPs demonstrated the presence of two clusters and deep divergence between Angola and all other individuals from Namibia and South Africa. These results provide evidence for adaptive divergence in response to a heterogeneous seascape in a large, mobile shark despite high levels of gene flow. These results are expected to inform management strategies and policy at the national and regional level for conservation of C. brachyurus populations.

Thermal plasticity in development and diapause strategy in a temperate butterfly across a latitudinal gradient

Abstract Trade‐offs among traits are central to life‐history theory and often closely linked to an organism's fitness. Understanding how these trade‐offs vary among populations and across environments is therefore important to more accurately predict species' responses to future climate change. However, the extent to which responses vary across populations remains unknown because few studies investigate intraspecific differences. We performed a full‐factorial split‐brood common garden experiment to test how variation in rearing temperature affects developmental timing and other traits important for survival during diapause in the Glanville fritillary butterfly (Melitaea cinxia). Pre‐diapause larvae originating from four regions across a latitudinal cline across Europe were reared at four temperatures (25, 28, 31 and 34°C), and we used a reaction norm approach to test for evidence of genetic differentiation and variation in developmental plasticity across regions. We found clear signs of genetic differentiation in multiple developmental traits, as well as differences in developmental plasticity. Northern larvae entered diapause in the fourth instar when the temperatures were low, whereas southern larvae did so in the fifth instar. As a result, development time is canalized with regards to temperature in northern larvae: due to entering diapause one stage earlier, they develop fast even in the cold, whereas southern larvae always develop slower, especially at low temperatures. As a trade‐off, northern larvae have a lower body mass when reared at cooler temperatures compared to southern larvae, and they show increased plasticity in diapause mass. No clear clinal patterns were found in relative fat content. Our results show that trade‐offs between body size, development time and growth rate can vary within species living across environmental clines, possibly as a consequence of natural selection to local environmental conditions or other genetic constraints. This variation highlights the importance of recognizing the context dependency of relationships between important life‐history traits and their interactions with local environments in predicting species' responses to climate change. Read the free Plain Language Summary for this article on the Journal blog.

Regional variation in climate change alters the range-wide distribution of colour polymorphism in a wild bird.

According to Gloger's rule, animal colouration is expected to be darker in wetter and warmer climates. Such environmental clines are predicted to occur in colour polymorphic species and to be shaped by selection if colour morphs represent adaptations to different environments. We studied if the distribution of the colour polymorphic tawny owl (Strix aluco) morphs (a pheomelanic brown and a pale grey) across Europe follow the predictions of Gloger's rule and if there is a temporal change in the geographical patterns corresponding to regional variations in climate change. We used data on tawny owl museum skin specimen collections. First, we investigated long-term spatiotemporal variation in the probability of observing the colour morphs in different climate zones. Second, we studied if the probability of observing the colour morphs was associated with general climatic conditions. Third, we studied if weather fluctuations prior to the finding year of an owl explain colour morph in each climate zone. The brown tawny owl morph was historically more common than the grey morph in every studied climate zone. Over time, the brown morph has become rarer in the temperate and Mediterranean zone, whereas it has first become rarer but then again more common in the boreal zone. Based on general climatic conditions, winter and summer temperatures were positively and negatively associated with the proportion of brown morph, respectively. Winter precipitation was negatively associated with the proportion of brown morph. The effects of 5-year means of weather on the probability to observe a brown morph differed between climate zones, indicating region-dependent effect of climate change and weather on tawny owl colouration. To conclude, tawny owl colouration does not explicitly follow Gloger's rule, implying a time and space-dependent complex system shaped by many factors. We provide novel insights into how the geographic distribution of pheomelanin-based colour polymorphism is changing.

Variation in gene expression along an elevation gradient of Rhododendron sanguineum var. haemaleum assessed in a comparative transcriptomic analysis.

Selection along environmental gradients may play a vital role in driving adaptive evolution. Nevertheless, genomic variation and genetic adaptation along environmental clines remains largely unknown in plants in alpine ecosystems. To close this knowledge gap, we assayed transcriptomic profiles of late flower bud and early leaf bud of Rhododendron sanguineum var. haemaleum from four different elevational belts between 3,000 m and 3,800 m in the Gaoligong Mountains. By comparing differences in gene expression of these samples, a gene co-expression network (WGCNA) was constructed to identify candidate genes related to elevation. We found that the overall gene expression patterns are organ-specific for the flower and leaf. Differentially expressed unigenes were identified in these organs. In flowers, these were mainly related to terpenoid metabolism (RsHMGR, RsTPS), while in leaves mainly related to anthocyanin biosynthesis (RsCHS, RsF3'5'H). Terpenoids are the main components of flower scent (fragrance) likely attracting insects for pollination. In response to fewer pollinators at higher elevation zone, it seems relatively less scent is produced in flower organs to reduce energy consumption. Secondary metabolites in leaves such as anthocyanins determine the plants' alternative adaptive strategy to extreme environments, such as selective pressures of insect herbivory from environmental changes and substrate competition in biosynthesis pathways at high elevations. Our findings indicated that the gene expression profiles generated from flower and leaf organs showed parallel expression shifts but with different functionality, suggesting the existence of flexibility in response strategies of plants exposed to heterogeneous environments across elevational gradients. The genes identified here are likely to be involved in the adaptation of the plants to these varying mountainous environments. This study thus contributes to our understanding of the molecular mechanisms of adaptation in response to environmental change.

Rapid diversification underlying the global dominance of a cosmopolitan phytoplankton

Marine phytoplankton play important roles in the global ecosystem, with a limited number of cosmopolitan keystone species driving their biomass. Recent studies have revealed that many of these phytoplankton are complexes composed of sibling species, but little is known about the evolutionary processes underlying their formation. Gephyrocapsa huxleyi, a widely distributed and abundant unicellular marine planktonic algae, produces calcified scales (coccoliths), thereby significantly affects global biogeochemical cycles via sequestration of inorganic carbon. This species is composed of morphotypes defined by differing degrees of coccolith calcification, the evolutionary ecology of which remains unclear. Here, we report an integrated morphological, ecological and genomic survey across globally distributed G. huxleyi strains to reconstruct evolutionary relationships between morphotypes in relation to their habitats. While G. huxleyi has been considered a single cosmopolitan species, our analyses demonstrate that it has evolved to comprise at least three distinct species, which led us to formally revise the taxonomy of the G. huxleyi complex. Moreover, the first speciation event occurred before the onset of the last interglacial period (~140 ka), while the second followed during this interglacial. Then, further rapid diversifications occurred during the most recent ice-sheet expansion of the last glacial period and established morphotypes as dominant populations across environmental clines. These results suggest that glacial-cycle dynamics contributed to the isolation of ocean basins and the segregations of oceans fronts as extrinsic drivers of micro-evolutionary radiations in extant marine phytoplankton.

Are populations less genetically diverse and more differentiated at the species range edges? Analysis of the quantitative trait and molecular variation in wild oat Avena sterilis

Abstract Although understanding factors determining the genetic makeup of natural populations has long been an important goal of evolutionary biology, the effect of population position within the species range (i.e., interior vs. edge) on species genetic variation is still unclear. According to the ‘abundant center’ hypothesis, the range edge populations are expected to exhibit lower genetic variation and higher differentiation than core populations because of their greater spatial isolation and smaller size. We tested these predictions by comparing the extent and structure of neutral (SSR) and quantitative trait genetic variation in 20 populations of an annual grass Avena sterilis hierarchically sampled at the species range core and two opposite edges. Within-population genetic diversity was reduced at the desert range edge compared to the range core as assessed by six SSR markers but not by eight quantitative traits; no reduction was detected at the mountain edge. Genetic differentiation among populations was higher at the desert range edge than the range core in both molecular markers and quantitative traits, but not at the mountain edge. Our results imply that the pattern of population genetic variation at the species range edges largely depends on the steepness of the environmental cline that has a major effect on species fitness. The more gradual the environmental cline from the species interior towards the edge, the higher the probability of detecting reduced genetic diversity and increased differentiation of peripheral populations as predicted by the ‘abundant center’ hypothesis.

Phylogeography and taxonomy of Coleonyx elegans Gray 1845 (Squamata: Eublepharidae) in Mesoamerica: The Isthmus of Tehuantepec as an environmental barrier

Population divergence leading to speciation is often explained by physical barriers causing allopatric distributions of historically connected populations. Environmental barriers have increasingly been shown to cause population divergence through local adaptation to distinct ecological characteristics. In this study, we evaluate population structuring and phylogeographic history within the Yucatán banded gecko Coleonyx elegans Gray 1845 to assess the role of both physical and environmental barriers in shaping the spatio-genetic distribution of a Mesoamerican tropical forest taxon. We generated RADseq and multi-locus Sanger datasets that included sampling across the entire species’ range. Results find support for two distinct evolutionary lineages that diverged during the late Pliocene and show recent population expansions. Furthermore, these genetic lineages largely align with subspecies boundaries defined by morphology. Several mountain ranges identified as phylogeographic barriers in other taxa act as physical barriers to gene flow between the two clades. Despite the absence of a physical barrier between lineages across the lowland Isthmus of Tehuantepec, no introgression was observed. Here, a steep environmental cline associated with seasonality of precipitation corresponds exactly with the distributional limits of the lineages, whose closest samples are only 30 km apart. The combination of molecular and environmental evidence, and in conjunction with previous morphological evidence, allows us to reassess the current taxonomy in an integrative framework. Based on our findings, we elevate the previously recognized subspecies from the Pacific versant, the Colima banded gecko C. nemoralis Klauber 1945, to full species status and comment on conservation implications.

Genetic Diversity and Structure of a Diverse Population of Picea sitchensis Using Genotyping-by-Sequencing

Picea sitchensis, Sitka spruce, is of interest to forestry as both a conservation species and a highly productive crop. Its native range stretches from Alaska to California, and it is hence distributed across a large environmental cline with areas of local adaptation. The IUFRO collection, established in 1968–1970, consists of 81 provenances of commercial and scientific interest spanning this native range. We used genotyping-by-sequencing on 1177 genotypes, originating from 80 of the IUFRO provenances which occupy 19 geographic regions of the Pacific Northwest, resulting in an SNP database of 36,567 markers. We detected low levels of genetic differentiation across this broad environmental cline, in agreement with other studies. However, we discovered island effects on geographically distant populations, such as those on Haida Gwaii and Kodiak Island. Using glaciation data, alongside this database, we see apparent post-glacial recolonization of the mainland from islands and the south of the range. Genotyping the IUFRO population expands upon the use of the collection in three ways: (i) providing information to breeders on genetic diversity which can be implemented into breeding programs, optimizing genetic gain for important traits; (ii) serving a scientific resource for studying spruce species; and (iii) utilizing provenances in breeding programs which are more tolerant to climate change.

Tension zone trapped by exogenous cline: Analysis of a narrow hybrid zone between two parapatric Oxytropis species (Fabaceae).

Hybrid zones have been widely highlighted for their interest in understanding evolutionary processes. It is generally accepted that hybrid zones can be maintained in a balance between dispersal and selection. However, the selective forces can either be endogenous (i.e., genetic incompatibilities between parental taxa) or exogenous (i.e., parental taxa are adapted to different environments). In this study, we evaluated these alternatives and determined the maintenance of a narrow hybrid zone between parapatric distributed Oxytropis diversifolia and O. leptophylla in Nei Mongol, China. For 507 individuals sampled from two populations in the hybrid zone, 12 O. diversifolia populations and five O. leptophylla populations, we measured leaf‐morphological characteristics, quantified genetic structure using 11 nuclear microsatellite loci and five chloroplast DNA intergenic regions, collected micro‐ and macrohabitat data, and conducted geographical cline analysis. We found that the two species differed in leaf morphology, and putative hybrids showed either intermediacy or a bias to O. diversifolia. Parental taxa formed two genetically distinct clusters, while populations in the hybrid zone consisted of both parental forms and various admixed individuals, exhibiting a bimodal pattern. The hybrid zone was coupled to ecological transitions of both microhabitat (i.e., the slope) and macroclimatic conditions. However, the genetic clines were significantly narrower than the environmental cline. Our results indicate that endogenous selection can be primarily responsible for maintaining the hybrid zone, while local adaptation accounts for the position of the zone. We further suggest the probable outcome of hybridization could be introgression.

Physiological characterization of Typhlatya cave shrimps: linking habitat with aerobic metabolism

The anchialine environment is characterized by a vertical stratification of water masses with different salinities. Cave shrimps of the genus Typhlatya are widespread inhabitants of the aquifer in fresh, brackish, and marine groundwater. Here we describe physiological aspects of three of the most abundant and widespread Typhlatya species that thrive in the fresh and marine groundwater habitats of the anchialine ecosystem of the Yucatan Peninsula. The aerobic scope (AS) of Typhlatya mitchelli, Typhlatya pearsei and Typhlatya dzilamensis was estimated through induced physical activity, whilst monitoring protein carbonylation and lipid peroxidation (as indicators of cellular damage), lactate accumulation (as an indicator of anaerobic metabolism) and the antioxidant system response. The critical thermal limits (CTL) of all three species as an additional measure of physiological plasticity were also determined. Our results showed that metabolic rates, AS and CTL were similar amongst the two species that inhabit fresh groundwater habitats, and differed markedly from T. dzilamensis, a species typically found in marine groundwater. The antioxidant system response in all three Typhlatya species accompanied the levels of aerobic metabolism following physical activity. However, the large amount of GSH observed in T. dzilamensis may be indicative of an adaptive trait to a more heterogeneous environment. The differences observed among Typhlatya species reflect different physiological adaptations that correspond to the environmental heterogeneity of their natural habitats. Our results suggest that the marine groundwater species, T. dzilamensis, could be better prepared to respond to a naturally more heterogeneous environment, in contrast to Typhlatya mitchelli and Typhlatya pearsei which rarely face environmental clines in the fresh groundwater habitat. Our findings contribute to a better understanding of the consequences of environmental change on ecologically important species that are restricted to live in the aquifer.

Functional evidence supports adaptive plant chemical defense along a geographical cline

Environmental clines in organismal defensive traits are usually attributed to stronger selection by enemies at lower latitudes or near the host's range center. Nonetheless, little functional evidence has supported this hypothesis, especially for coevolving plants and herbivores. We quantified cardenolide toxins in seeds of 24 populations of common milkweed (Asclepias syriaca) across 13 degrees of latitude, revealing a pattern of increasing cardenolide concentrations toward the host's range center. The unusual nitrogen-containing cardenolide labriformin was an exception and peaked at higher latitudes. Milkweed seeds are eaten by specialist lygaeid bugs that are even more tolerant of cardenolides than the monarch butterfly, concentrating most cardenolides (but not labriformin) from seeds into their bodies. Accordingly, whether cardenolides defend seeds against these specialist bugs is unclear. We demonstrate that Oncopeltus fasciatus (Lygaeidae) metabolized two major compounds (glycosylated aspecioside and labriformin) into distinct products that were sequestered without impairing growth. We next tested several isolated cardenolides invitro on the physiological target of cardenolides (Na+/K+-ATPase); there was little variation among compounds in inhibition of an unadapted Na+/K+-ATPase, but tremendous variation in impacts on that of monarchs and Oncopeltus. Labriformin was the most inhibitive compound tested for both insects, but Oncopeltus had the greater advantage over monarchs in tolerating labriformin compared to other compounds. Three metabolized (and stored) cardenolides were less toxic than their parent compounds found in seeds. Our results suggest that a potent plant defense is evolving by natural selection along a geographical cline and targets specialist herbivores, but is met by insect tolerance, detoxification, and sequestration.