Plant resource-use strategies: the importance of phenotypic plasticity in response to a productivity gradient for two subalpine species

Background and Aims

Functional traits are indicators of plant interactions with their environment and the resource-use strategies of species can be defined through some key functional traits. The importance of genetic variability and phenotypic plasticity in trait variations in response to a common environmental change was investigated in two subalpine species.

Methods

Two species with contrasted resource-use strategies, Dactylis glomerata and Festuca paniculata, were grown along a productivity gradient in a greenhouse experiment. Functional traits of different genotypes were measured to estimate the relative roles of phenotypic plasticity and genetic variability, and to compare their levels of phenotypic plasticity.

Key Results

Trait variability in the field for the two species is more likely to be the result of phenotypic plasticity rather than of genetic differentiation between populations. The exploitative species D. glomerata expressed an overall higher level of phenotypic plasticity compared with the conservative species F. paniculata. In addition to different amplitudes of phenotypic plasticity, the two species differed in their pattern of response for three functional traits relevant to resource use (specific leaf area, leaf dry matter content and leaf nitrogen content).

Conclusions

Functional trait variability was mainly the result of phenotypic plasticity, with the exploitative species showing greater variability. In addition to average trait values, two species with different resource-use strategies differed in their plastic responses to productivity.     

Plasticity of functional traits varies clinally along a rainfall gradient in Eucalyptus tricarpa

Widespread species often occur across a range of climatic conditions, through a combination of local genetic adaptations and phenotypic plasticity. Species with greater phenotypic plasticity are likely to be better positioned to cope with rapid anthropogenic climate changes, while those displaying strong local adaptations might benefit from translocations to assist the movement of adaptive genes as the climate changes. Eucalyptus tricarpa occurs across a climatic gradient in south‐eastern Australia, a region of increasing aridity, and we hypothesized that this species would display local adaptation to climate. We measured morphological and physiological traits reflecting climate responses in nine provenances from sites of 460 to 1040 mm annual rainfall, in their natural habitat and in common gardens near each end of the gradient. Local adaptation was evident in functional traits and differential growth rates in the common gardens. Some traits displayed complex combinations of plasticity and genetic divergence among provenances, including clinal variation in plasticity itself. Provenances from drier locations were more plastic in leaf thickness, whereas leaf size was more plastic in provenances from higher rainfall locations. Leaf density and stomatal physiology (as indicated by δ¹³C and δ¹⁸O) were highly and uniformly plastic. In addition to variation in mean trait values, genetic variation in trait plasticity may play a role in climate adaptation.     

Trait plasticity,not values,best corresponds with woodland plant success in novel and manipulated habitats

Aims The clustering of plants with similar leaf traits along environmental gradients may arise from adaptation as well as acclimation to heterogeneous habitat conditions. Determining the forces that shape plant leaf traits requires both linking variation in trait morphology with abiotic gradients and linking that trait variation with plant performance under varying abiotic conditions. Across the spectrum of plant types, shade-tolerant evergreen herbs are relatively low in trait plasticity, compared to deciduous and sun-adapted species. These plants employ stress-tolerant strategies for survival, which coincide with relatively static trait morphologies, slow growth and hence a lower ability to adjust to changing environmental conditions.Methods We investigate how the survival of two ecologically similar understory evergreen species, Asarum arifolium and Hepatica nobilis, corresponds with variation in six commonly measured functional traits (leaf area, specific leaf area, plant height, leaf number, leaf length and shoot mass) along natural and experimental abiotic gradients. We examine temporal (the period 2007–9) and spatial (100 km) variations in these traits after (i) translocating 576 plants across a span from the southern Appalachian Mountains in NC, USA, to the Piedmont, GA, USA, which includes north- and south-facing slope habitats and (ii) the experimental manipulation of diffuse light and soil moisture.Important findings We find that when translocated into a novel habitats, with novel environmental conditions that often are more extreme than the source habitat, both species appear capable of considerable morphological acclimation and generally converge to similar trait values. Hepatica nobilis does not exhibit mean trait values particularly different from those of A. arifolium, but it demonstrates much greater phenotypic plasticity. These results indicate that relatively conservative plant species nonetheless acclimate and survive across heterogeneous environmental conditions.     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

Phenotypic plasticity closely linked to climate at origin and resulting in increased mortality under warming and frost stress in a common grass

Phenotypic plasticity is important for species responses to global change and species coexistence. Phenotypic plasticity differs among species and traits and changes across environments. Here, we investigated phenotypic plasticity of the widespread grass Arrhenatherum elatius in response to winter warming and frost stress by comparing phenotypic plasticity of 11 geographically and environmentally distinct populations of this species to phenotypic plasticity of populations of different species originating from a single environment. The variation in phenotypic plasticity was similar for populations of a single species from different locations compared to populations of functionally and taxonomically diverse species from one environment for the studied traits (leaf biomass production and root integrity after frost) across three indices of phenotypic plasticity (RDPI, PIN, slope of reaction norm). Phenotypic plasticity was not associated with neutral genetic diversity but closely linked to the climate of the populations’ origin. Populations originating from warmer and more variable climates showed higher phenotypic plasticity. This indicates that phenotypic plasticity can itself be considered as a trait subject to local adaptation to climate. Finally, our data emphasize that high phenotypic plasticity is not per se positive for adaptation to climate change, as differences in stress responses are resulting in high phenotypic plasticity as expressed by common plasticity indices, which is likely to be related to increased mortality under stress in more plastic populations.     

Differences in the leaf functional traits of six beech (Fagus sylvatica L.) populations are reflected in their response to water limitation

Patterns of intraspecific variation in functional traits have been widely studied across plant species to find out what general suites of traits provide functional advantage under specific environmental conditions. Much less is known about this variation within tree species and, in particular, about its relationship with performance variables such as photosynthetic rates under water deficit. Nevertheless, this knowledge is fundamental to understand the adaptive potential of drought sensitive tree species to increased aridity as predicted in the context of climate change.Intraspecific variation in photosynthetic performance and other leaf functional traits in response to water availability were examined in a glasshouse experiment using seedlings of six European beech populations. The physiological response of seedlings to a “water stress” treatment was compared to a “control” treatment along an experimental cycle of progressive soil water deficit and recovery. We found evidence of intraspecific variation in beech's photosynthetic performance and other leaf functional traits in response to water availability. We also detected intraspecific variation in leaf-level tolerance of water deficit and phenotypic plasticity to water availability suggesting a pattern shaped by both regional and local scale effects. The Swedish population was particularly sensitive to water deficit, being the only population showing impaired photochemical efficiency under the experimental water deficit. Leaf-level tolerance of water deficit was related to PNUE, but not to other functional traits, such as WUE, SLA or leaf nitrogen content, that have been described to vary across species in adaptation to drought tolerance. Our results support the idea that general trends for variation in functional traits across species do not necessarily reflect a similar pattern when observed at the intraspecific level. The observed functional variation between beech populations reaffirms the importance of local adaptation to water deficit in the context of climate change.     

Phenotypic plasticity and population differentiation in response to salinity in the invasive cordgrass <Emphasis Type="Italic">Spartina densiflora</Emphasis>

Salinity and tidal inundation induce physiological stress in vascular plant species and influence their distribution and productivity in estuarine wetlands. Climate change-induced sea level rise may magnify these abiotic stressors and the physiological stresses they can cause. Understanding the potential of invasive plants to respond to predicted salinity increases will elucidate their potential niche breadth. To examine potential phenotypic plasticity and functional trait responses to salinity stress in the invasive cordgrass Spartina densiflora, we collected rhizomes from four invasive populations occurring from California to Vancouver Island, British Columbia on the Pacific Coast of North America. In a glasshouse common garden experiment, we measured plant traits associated with growth and allocation, photosynthesis, leaf pigments, and leaf chemistry and calculated plasticity indices across imposed salinity treatments. Fifteen of 21 leaf chemistry, pigment, morphological and physiological traits expressed plastic responses to salinity. When averaged across all measured traits, degree of plasticity did not vary among sampled populations. However, differences in plasticity among populations in response to salinity were observed for 9 of 21 measured plant traits. Leaf chemistry and adaxial leaf rolling trait responses demonstrated the highest degree of plasticity, while growth and allocation measures were less plastic. Phenotypic plasticity of leaf functional traits to salinity indicates the potential of S. densiflora to maintain invasive growth in response to rising estuarine salinity with climate change.     

The plasticity of phenotypic integration in response to light and water availability in the pepper grass, <Emphasis Type="Italic">Lepidium bonariense</Emphasis>

Organisms that live in a heterogeneous environment face a number of important challenges. On one hand, they require the flexibility to respond to environmental conditions and change their phenotype accordingly. On the other, they are required to be robust in their overall body plan to ensure an integrated, functional organism. Here, we examine the relationship between phenotypic plasticity and integration in the common peppergrass, Lepidium bonariense, by examining the multivariate response of a series of functional traits to a combination of light and water treatments. Lepidium bonariense displayed considerable variation in phenotype in response to water and light availability with the extraction of the first two principal components retaining 85% of the total variation in our data set. Principal component 1 (PC1) largely reflects the negative genetic correlation between specific leaf area and overall plant size, whereas PC2 was typical of a shade-avoidance syndrome displayed by many species of vascular plants. Both PC1 and PC2 exhibited considerable variation among genotypes in phenotypic plasticity in response to the combined effect of light and water availability. Despite complex plasticity in this species, we demonstrate that variation in light and water availability did not significantly influence patterns of functional trait integration, with the genetic variance–covariance matrix remaining stable across environments.     

Light-dependent leaf trait variation in 43 tropical dry forest tree species

Our understanding of leaf acclimation in relation to irradiance of fully grown or juvenile trees is mainly based on research involving tropical wet forest species. We studied sun-shade plasticity of 24 leaf traits of 43 tree species in a Bolivian dry deciduous forest. Sampling was confined to small trees. For each species, leaves were taken from five of the most and five of the least illuminated crowns. Trees were selected based on the percentage of the hemisphere uncovered by other crowns. We examined leaf trait variation and the relation between trait plasticity and light demand, maximum adult stature, and ontogenetic changes in crown exposure of the species. Leaf trait variation was mainly related to differences among species and to a minor extent to differences in light availability. Traits related to the palisade layer, thickness of the outer cell wall, and N(area) and P(area) had the greatest plasticity, suggesting their importance for leaf function in different light environments. Short-lived pioneers had the highest trait plasticity. Overall plasticity was modest and rarely associated with juvenile light requirements, adult stature, or ontogenetic changes in crown exposure. Dry forest tree species had a lower light-related plasticity than wet forest species, probably because wet forests cast deeper shade. In dry forests light availability may be less limiting, and low water availability may constrain leaf trait plasticity in response to irradiance.     

Geographic variation and plasticity to water and nutrients in Pelargonium australe

Here, patterns of phenotypic plasticity and trait integration of leaf characteristics in six geographically discrete populations of the perennial herb Pelargonium australe were compared. It was hypothesized that populations would show local adaptation in trait means, but similar patterns of plasticity and trait integration. Further, it was questioned whether phenotypic plasticity was positively correlated with environmental heterogeneity and whether plasticity for water-use traits in particular was adaptive. Seedlings were grown in a glasshouse at six combinations of water and nutrient availability. Leaf anatomical, morphological and gas exchange traits were measured. High amounts of plasticity in leaf traits were found in response to changes in growth conditions and there was evidence of local adaptation among the populations. While there were significant correlations between plasticity and environmental heterogeneity, not all were positive. Notably, patterns of plasticity and trait integration varied significantly among populations. Despite that variation, some of the observed plasticity was adaptive: fitness was correlated with conservative water use when water was limiting. Pelargonium arrived in Australia approximately 5 million yr ago. It is concluded here that high amounts of plasticity, in some cases adaptive, and weak integration among traits may be key to the spread and success of this species.     

Hummingbird blood traits track oxygen availability across space and time

Predictable trait variation across environments suggests shared adaptive responses via repeated genetic evolution, phenotypic plasticity or both. Matching of trait–environment associations at phylogenetic and individual scales implies consistency between these processes. Alternatively, mismatch implies that evolutionary divergence has changed the rules of trait–environment covariation. Here we tested whether species adaptation alters elevational variation in blood traits. We measured blood for 1217 Andean hummingbirds of 77 species across a 4600-m elevational gradient. Unexpectedly, elevational variation in haemoglobin concentration ([Hb]) was scale independent, suggesting that physics of gas exchange, rather than species differences, determines responses to changing oxygen pressure. However, mechanisms of [Hb] adjustment did show signals of species adaptation: Species at either low or high elevations adjusted cell size, whereas species at mid-elevations adjusted cell number. This elevational variation in red blood cell number versus size suggests that genetic adaptation to high altitude has changed how these traits respond to shifts in oxygen availability.     

Intraspecific trait variation across elevation predicts a widespread tree species' climate niche and range limits

Global change is widely altering environmental conditions which makes accurately predicting species range limits across natural landscapes critical for conservation and management decisions. If climate pressures along elevation gradients influence the distribution of phenotypic and genetic variation of plant functional traits, then such trait variation may be informative of the selective mechanisms and adaptations that help define climatic niche limits. Using extensive field surveys along 16 elevation transects and a large common garden experiment, we tested whether functional trait variation could predict the climatic niche of a widespread tree species (Populus angustifolia) with a double quantile regression approach. We show that intraspecific variation in plant size, growth, and leaf morphology corresponds with the species' total climate range and certain climatic limits related to temperature and moisture extremes. Moreover, we find evidence of genetic clines and phenotypic plasticity at environmental boundaries, which we use to create geographic predictions of trait variation and maximum values due to climatic constraints across the western US. Overall, our findings show the utility of double quantile regressions for connecting species distributions and climate gradients through trait‐based mechanisms. We highlight how new approaches like ours that incorporate genetic variation in functional traits and their response to climate gradients will lead to a better understanding of plant distributions as well as identifying populations anticipated to be maladapted to future environments.     

格氏栲天然林林窗植物群落功能性状的变异

林窗是森林更新演替的重要环节, 揭示林窗环境下功能性状变异来源及其相对贡献, 有助于阐明植物对林窗环境的响应。该研究以中亚热带格氏栲(Castanopsis kawakamii)天然林为对象, 设置9个不同大小的林窗样地, 运用方差分解探讨林窗、物种和个体对叶性状变异的相对贡献, 采用线性回归分析不同大小林窗下群落性状变化及种间和种内性状变异的重要性。研究发现: (1)格氏栲天然林林窗植物比叶面积、叶干物质含量、叶厚和叶绿素含量由种间性状变异主导, 叶氮含量由种内性状变异主导, 叶磷含量受林窗大小影响最大。(2)群落叶磷含量与林窗大小具有显著正相关关系, 土壤温度和水解氮含量对群落叶磷含量具有显著正效应, 土壤有效磷含量具有显著负效应。(3)沿林冠开放度的群落叶磷含量变化主要由种内性状变异引起, 优势种扮演着重要角色。结果表明, 格氏栲天然林林窗环境下植物功能性状仍以种间性状变异为主(平均41%), 但沿林窗环境梯度的群落性状变化主要源自种内性状变异, 通过植物表型可塑性响应环境改变, 优势种作用明显。     

桂林岩溶石山青冈群落植物功能性状的尺度变化与关联

研究植物功能性状在不同尺度的变异和关联,对于揭示植物对环境的适应策略和群落构建规律具有重要意义。以岩溶石山青冈群落为研究对象,测量了研究区内20个样方74种木本植物的叶面积、比叶面积和木材密度3个功能性状值,利用性状梯度分析法分析了3个性状在群落内部(α组分)及群落间(β组分)的变异格局及相关性。结果表明:(1)群落内3个植物功能性状的α值范围均大于β值范围,即物种相对于共生物种性状值的变化大于沿着群落平均性状梯度的变化。(2)植物功能性状比叶面积的种内差异引起的变化小于群落水平。(3)叶面积与比叶面积、比叶面积与木材密度、叶面积与木材密度的β组分相关性均最强,而α组分间无相关性或相关性较弱,即叶面积与比叶面积、比叶面积与木材密度、叶面积与木材密度两两性状间的相关性在群落间的依赖程度比群落内共生物种的依赖性要强,暗示物种在群落内和群落间采取不同的生态策略来适应环境。     

Effect of cold and menthol on membrane potential in plants

'The Holy Grail' of plant ecology is to uncover rules that associate species and traits with environmental constraints, community composition and subsequent ecosystem functioning. These aims have been crystallized in recent years within the context of global climate change and environmental pollution, increasing the urgency of the need to predict how vegetation will respond across spatial scales. We investigated whether genetic diversity is associated with the way in which phenotypic plasticity within plant populations is realized and whether this is related to genotype abundance. We used environmental metabolomics to demonstrate biochemical variation between co-occurring genotypes of Carex caryophyllea L. A novel combined metabolomic/functional trait analysis was used to test the functionality of this variation in governing plasticity to variation in edaphic conditions, with particular reference to metabolic pathways that play important roles in growth-related traits. We show that genetic diversity within a wild C. caryophyllea population relates to differences in metabolic composition and functional traits in response to soil nutrient variation, influencing genotype abundance within a community. Our findings highlight the vital role genetic diversity plays within a population in facilitating plant phenotypic plasticity and the potential usefulness of environmental metabolomics to future ecological studies.     

Is seasonal variation in leaf traits adaptive for the annual plant Dicerandra linearifolia?

Empirical studies of phenotypic plasticity have often relied on the plausibility that a plastic response to the environment would increase fitness in order to diagnose the response as adaptive. I conducted a test of the hypothesis that seasonal variation in leaf traits is an adaptive response to seasonal variation in environmental conditions faced by the annual plant Dicerandralinearifolia. This species exhibits variation in leaf morphology and anatomy in response to temperature that is consistent with the expectations for adaptive plasticity. I examined variation in the size, thickness and density of stomata of leaves that develop in summer and winter and used analysis of phenotypic selection during winter and summer seasons to test the hypothesis that seasonal variation in these traits is adaptive. Regression analyses of estimated dry mass (as a proxy for fitness) on leaf traits revealed no evidence supporting the adaptive hypothesis. Selection favoured individuals with large and thick leaves in both winter and summer, and density of stomata had little or no effect on estimated relative fitness in any season. Correspondence between seasonal variation in leaf thickness and density of stomata and expectations for adaptive plasticity appears to be purely fortuitous. Seasonal variation in leaf traits may persist simply because there is no selection against individuals in which these traits vary. My results underscore the importance of definitive tests of the hypothesis of adaptation to distinguish adaptive plasticity from neutral or nonadaptive phenotypic plasticity.     

Intraspecific Trait Variation Driven by Plasticity and Ontogeny in Hypochaeris radicata

The importance of intraspecific variation in plant functional traits for structuring communities and driving ecosystem processes is increasingly recognized, but mechanisms governing this variation are less studied. Variation could be due to adaptation to local conditions, plasticity in observed traits, or ontogeny. We investigated 1) whether abiotic stress caused individuals, maternal lines, and populations to exhibit trait convergence, 2) whether trait variation was primarily due to ecotypic differences or trait plasticity, and 3) whether traits varied with ontogeny. We sampled three populations of Hypochaeris radicata that differed significantly in rosette diameter and specific leaf area (SLA). We grew nine maternal lines from each population (27 lines total) under three greenhouse conditions: ambient conditions (control), 50% drought, or 80% shade. Plant diameter and relative chlorophyll content were measured throughout the experiment, and leaf shape, root∶shoot ratio, and SLA were measured after five weeks. We used hierarchical mixed-models and variance component analysis to quantify differences in treatment effects and the contributions of population of origin and maternal line to observed variation. Observed variation in plant traits was driven primarily by plasticity. Shade significantly influenced all measured traits. Plant diameter was the only trait that had a sizable proportion of trait variation (30%) explained by population of origin. There were significant ontogenetic differences for both plant diameter and relative chlorophyll content. When subjected to abiotic stress in the form of light or water limitation, Hypochaeris radicata exhibited significant trait variability. This variation was due primarily to trait plasticity, rather than to adaptation to local conditions, and also differed with ontogeny.     

Potential local adaptation in populations of invasive reed canary grass (Phalaris arundinacea) across an urbanization gradient

Urban stressors represent strong selective gradients that can elicit evolutionary change, especially in non‐native species that may harbor substantial within‐population variability. To test whether urban stressors drive phenotypic differentiation and influence local adaptation, we compared stress responses of populations of a ubiquitous invader, reed canary grass (Phalaris arundinacea). Specifically, we quantified responses to salt, copper, and zinc additions by reed canary grass collected from four populations spanning an urbanization gradient (natural, rural, moderate urban, and intense urban). We measured ten phenotypic traits and trait plasticities, because reed canary grass is known to be highly plastic and because plasticity may enhance invasion success. We tested the following hypotheses: (a) Source populations vary systematically in their stress response, with the intense urban population least sensitive and the natural population most sensitive, and (b) plastic responses are adaptive under stressful conditions. We found clear trait variation among populations, with the greatest divergence in traits and trait plasticities between the natural and intense urban populations. The intense urban population showed stress tolerator characteristics for resource acquisition traits including leaf dry matter content and specific root length. Trait plasticity varied among populations for over half the traits measured, highlighting that plasticity differences were as common as trait differences. Plasticity in root mass ratio and specific root length were adaptive in some contexts, suggesting that natural selection by anthropogenic stressors may have contributed to root trait differences. Reed canary grass populations in highly urbanized wetlands may therefore be evolving enhanced tolerance to urban stressors, suggesting a mechanism by which invasive species may proliferate across urban wetland systems generally.     

Intraspecific trait variation and allocation strategies of calcareous grassland species: Results from a restoration experiment

Intra- and interspecific trait variation express the response of plants dealing with different environmental conditions. We measured root and leaf traits on 14 species of calcareous grasslands in a restoration experiment. We aimed at identifying intraspecific differences in biomass allocation and functional plant traits under contrasting soil conditions by comparing plants growing in ancient grassland and two restored grasslands on ex-arable land, one of them with topsoil removal. Relative importance of trait variation within and among species, and among site was assessed by variance partitioning. Interspecific variation was more important than intraspecific variation, but the contribution of the latter to total variation was considerable, especially for specific leaf area. Changes in soil properties due to topsoil removal resulted in lower values of plant height, specific leaf area and specific root length compared to the control (ancient grassland). Soil fertility found in the treatment without top soil removal did not affect plant plasticity compared to the control. The study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species. We conclude that caution must be taken when using mean trait values for plastic species or when working with environmental gradients.     

Drought affects the coordination of belowground and aboveground resource‐related traits in Solidago canadensis in China

Quantifying patterns of variation and coordination of plant functional traits can help to understand the mechanisms underlying both invasiveness and adaptation of plants. Little is known about the coordinated variations of performance and functional traits of different organs in invasive plants, especially in response to their adaptation to environmental stressors. To identify the responses of the invasive species Solidago canadensis to drought, 180 individuals were randomly collected from 15 populations and 212 ramets were replanted in a greenhouse to investigate both the response and coordination between root and leaf functional traits. Drought significantly decreased plant growth and most of the root and leaf functional traits, that is, root length, surface area, volume and leaf size, number, and mass fraction, except for the root length ratio and root mass fraction. Phenotypic plasticity was higher in root traits than in leaf traits in response to drought, and populations did not differ significantly. The plasticity of most root functional traits, that is, root length (RL), root surface area (RSA), root volume (RV), and root mass fraction (RMF), were significantly positively correlated with biomass between control and drought. However, the opposite was found for leaf functional traits, that is, specific leaf area (SLA), leaf area ratio (LAR), and leaf mass fraction (LMF). Drought enhanced the relationship between root and leaf, that is, 26 pairwise root–leaf traits were significantly correlated under drought, while only 15 pairwise root–leaf traits were significantly correlated under control conditions. Significant correlations were found between biomass and all measured functional traits except for leaf size. RV, root length ratio, RMF, total area of leaves, and LMF responded differently to water availability. These responses enable S. canadensis to cope with drought conditions and may help to explain the reason of the vast ecological amplitude of this species.