Plasticity Of Functional Traits Research Articles

Across-stage consequences of thermal stress have trait-specific effects and limited fitness costs in the harlequin ladybird, Harmonia axyridis

The fitness consequences of thermal plasticity of functional traits of insects, and the life stages at which these responses manifest, remain a subject of much debate. Here, we examine whether a short-term exposure to cold, medium or warm fluctuating temperature regimes during development (larval or adult) affects adult performance, thermal tolerance and fitness of the globally invasive harlequin ladybird Harmonia axyridis. We found plastic responses of a metric of heat stress resistance—critical thermal maximum—and of preferred body temperature after adult temperature exposure, but not in other traits measured. By contrast, exposure of larval stages resulted in plasticity of adult walking performance (height and breadth of the curve) but not tolerance or preference. We found distinct fitness responses between larval and adult treatments, but a composite fitness index revealed negligible effects on reproductive output. These results suggest that the drivers underlying the plasticity of temperature tolerance and temperature selection are different to those shaping the plasticity of walking speed. By testing specific predictions based on current theory of developmental and reversible plasticity, this study contributes novel data to plastic responses of behaviour, stress resistance and fitness to temperature exposure across life stages and thus, provides insights to the broader evolutionary and ecological significance of these responses.

Functional Trait Plasticity but Not Coordination Differs in Absorptive and Transport Fine Roots in Response to Soil Depth

Absorptive and transport fine roots (diameter ≤ 2 mm) differ greatly in anatomy, morphology, and physiology, as well as their responses to environmental changes. However, it is still not well understood how their functional traits and biomass repartition respond to resource variability associated with increasing soil depth. Herein, we sampled the first five order roots of three hardwoods, i.e., Juglans mandshurica Maxim., Fraxinus mandshurica Rupr., and Phellodendron amurense Rupr. at surface (0–10 cm) and subsurface (20–30 cm) soil layers, respectively, and measured root biomass, anatomy, morphology, chemistry, and physiology at the branch-order level. Based on the anatomical characteristics, absorptive and transport fine roots were identified within each order, and their amounts and functional trait plasticity to soil depth were examined. The results showed that across soil layers, the first three order roots were mainly absorptive roots, while the fourth- and fifth-order roots were transport ones. From surface to subsurface soil layers, both the number and biomass proportion of absorptive fine roots decreased but those of transport fine roots increased. Transport fine root traits were more plastic to soil depth than absorptive ones, especially for the conduit-related traits. Absorptive fine roots in surface soil generally had stronger potential for resource acquisition than those in deeper soil, as indicated by their longer specific root length and greater root branching density. In comparison, transport fine roots in deeper soil were generally enhanced in their transportation function, with wider stele and higher hydraulic conductivity. Our findings suggest that functional specialization via multi-trait plasticity and coordination in both absorptive and transport fine roots along the soil depth would benefit the efficient soil resource exploitation of trees in forest ecosystems.

Comparison of Leaf Functional Trait Plasticity between Exotic and Native <i>Ligustrum</i> Species in Japan

Comparison of Leaf Functional Trait Plasticity between Exotic and Native <i>Ligustrum</i> Species in Japan

Plasticity of functional traits and optimality of biomass allocation in elevational ecotypes of Arabidopsis halleri grown at different soil nutrient availabilities.

In mountainous areas, plant distribution is constrained by various environmental stresses. Plasticity and constancy in plant functional traits may relate to optimal strategies at respective habitats and to ecotypic differentiation along elevation. Although plant biomass allocation has been extensively studied in relation to adaptation to soil nutrient availability along elevation, its optimality is still poorly understood. We examined soil nutrient availability in the field and conducted growth analysis for two elevational ecotypes of Arabidopsis halleri grown under different nutrient availabilities. We determined plasticity in morphological and physiological traits and evaluated optimal biomass allocation using an optimality model. Our field investigation indicated that soil nitrogen (N) availability increased rather than decreased with increasing elevation. Our growth analysis revealed that lowland ecotype was more plastic in morphological variables and N concentrations, whereas the highland ecotype was more plastic in other physiological variables such as the net assimilation rate (NAR). The leaf mass ratio (LMR) in the lowland ecotype was moderately plastic at the whole range of N availabilities, whereas LMR in the highland ecotype was very plastic at higher N availabilities only. The optimality model indicated that the LMR of the lowland ecotype was nearly optimal throughout the range of studied N availabilities, whereas that of the highland ecotype was suboptimal at low N availability. These results suggest that highland ecotype is adapted only to high N availability, whereas the lowland ecotype is adapted to a relatively wide range of N availabilities as a result of natural selection in their respective habitats. We conclude that an adaptive differentiation has occurred between the two ecotypes and plasticity in the biomass allocation is directly related to its optimization in changing environments.

Plasticity of functional traits of tree of heaven is higher in exotic than in native habitats

Novel features related to turgor loss adjustment and higher ability to modulate leaf-construction costs could improve A. altissima fitness in its invaded range. Phenotypic plasticity has been recently proposed to play an important role in invasion process. More precisely, phenotypic plasticity of alien invasive species could be higher in their exotic rather than native habitat, probably because of the release from biotic or abiotic constraints. In this paper, several plant functional traits were analysed on Ailanthus altissima (Mill.) Swingle, a highly invasive species in Europe, aiming at providing a comparison among key functional traits measured on individuals growing in their native habitat and to test if values of functional traits and their related plasticity are higher in exotic habitats. Our analysis pinpointed that variability of functional traits in the native habitat was mainly driven by different irradiance, temperature and evaporative demand of the studied sites, in accordance to the traits trade-offs in the Leaf Economic Spectrum. Physiological traits related to drought tolerance were different between native and exotic habitats. In the native one, A. altissima relied on osmoregulation processes to adjust leaf turgor loss point (Ψtlp), as the osmotic potential at full turgor (π0) were lower in drier sites. In the exotic habitat, individuals in drier sites had similar π0 but lower wall elasticity (e), suggesting that leaves had thinner cell walls. Moreover, plasticity in specific leaf area and e were higher in the exotic habitat. The novel features related to Ψtlp adjustment and the higher plasticity in traits related to leaf-construction costs could increase A. altissima fitness in response to different environmental conditions in its invasive range.

Plastic biomass allocation as a trait increasing the invasiveness of annual bluegrass (Poa annua L.) in Antarctica

The plasticity of functional traits promotes invasiveness of a species. Biomass allocation, as one of these traits, is responsible for resource acquisition, and its plastic modifications can be of adaptive value in new environments before any genetic adaptations may occur. Our aim was to compare in situ biomass allocation in aboveground and belowground organs in an Antarctic and a Polish population of annual bluegrass (Poa annua), the only alien plant species successfully invading Antarctica. The Antarctic population was characterised by three times lower aboveground biomass, more compact plant growth habit and higher fraction of biomass allocated into belowground organs than in the Polish population. The differences between populations are probably a result of adaptation to local conditions. The modifications of the studied traits in the Antarctic population are most likely a response to extreme atmospheric and edaphic conditions and enable the species to survive and spread in this hostile environment. Our results are in accordance with the balanced growth hypothesis. At the same time, these trait values enhance species performance under Antarctic conditions making P. annua a potential threat to local plant communities under altering climate changes and growing human impact scenario.

Phenotypic plasticity and specialization along an altitudinal gradient in Trifolium repens

Phenotypic plasticity and specialization represent seemingly contrasting strategies to heterogeneous environments. Specialization is associated with the loss of phenotypic plasticity, particularly in functional traits. However, it is equivocal if this loss of plasticity is observed only in the specific habitat and stress type where the specialization occurs or a general loss of plasticity is seen across habitats. We examined populations of Trifolium repens L. following an expansion during the colonization of Australia from relatively good low altitude habitats to more stressful high altitude habitats in the Blue Mountains region, New South Wales, Australia. We examined if specialization to abiotic stress causes a loss of adaptive plasticity in functional traits under competition treatments (a different type of stress rather than abiotic stress). We found that both low and high altitude populations experienced a loss in performance in competition treatments and did not express a shade avoidance response under competition. Specialization to higher abiotic stress was associated with the loss of adaptive plasticity in functional traits. Our results suggest that specialization may limit the responses of plants to future environmental changes.

Above- and below-ground allocation and functional trait response to soil water inputs and drying rates of two common savanna grasses

The study of functional trait plasticity and optimal allocation strategies in a water competition context may help to explain the mechanisms relating plant competition to ecological semi-arid patterns. We measured four functional plant traits –root to shoot ratio, superficial to deep roots ratio, root diameter, and root estimated surface area to shoot ratio– in wild populations of two C4 grasses in a Kenyan savanna to assess their responses to different water inputs (changing with latitude) and soil drying rates (driven by canopy cover). Root:shoot allocation was different between the two species. We show that to account for root architecture helps understanding root allocation strategies, especially in species with high plasticity in root geometry. Concretely, in our case, a higher root allocation in wetter conditions was driven by water storage and not water foraging in the species with high root diameter variability. Superficial root allocation responded to changes associated with drying rates but not to water input, and shallower root systems developed under higher water stress conditions, supporting rainfall intermittency models of savanna tree-grass competition. We also suggest self-shading as a facilitation mechanism that may affect spatial patterns according to the scale-dependent feedback hypothesis.

Environment and host as large-scale controls of ectomycorrhizal fungi.

Explaining the large-scale diversity of soil organisms that drive biogeochemical processes-and their responses to environmental change-is critical. However, identifying consistent drivers of belowground diversity and abundance for some soil organisms at large spatial scales remains problematic. Here we investigate a major guild, the ectomycorrhizal fungi, across European forests at a spatial scale and resolution that is-to our knowledge-unprecedented, to explore key biotic and abiotic predictors of ectomycorrhizal diversity and to identify dominant responses and thresholds for change across complex environmental gradients. We show the effect of 38 host, environment, climate and geographical variables on ectomycorrhizal diversity, and define thresholds of community change for key variables. We quantify host specificity and reveal plasticity in functional traits involved in soil foraging across gradients. We conclude that environmental and host factors explain most of the variation in ectomycorrhizal diversity, that the environmental thresholds used as major ecosystem assessment tools need adjustment and that the importance of belowground specificity and plasticity has previously been underappreciated.

Linking plant functional trait plasticity and the large increase in forest water use efficiency

AbstractElevated atmospheric CO2 concentrations are expected to enhance photosynthesis and reduce stomatal conductance, thus increasing plant water use efficiency. A recent study based on eddy covariance flux observations from Northern Hemisphere forests showed a large increase in inherent water use efficiency (IWUE). Here we used an updated version of the same data set and robust uncertainty quantification to revisit these contemporary IWUE trends. We tested the hypothesis that the observed IWUE increase could be attributed to interannual trends in plant functional traits, potentially triggered by environmental change. We found that IWUE increased by ~1.3% yr−1, which is less than previously reported but still larger than theoretical expectations. Numerical simulations with the Tethys‐Chloris ecosystem model using temporally static plant functional traits cannot explain this increase. Simulations with plant functional trait plasticity, i.e., temporal changes in model parameters such as specific leaf area and maximum Rubisco capacity, match the observed trends in IWUE. Our results show that trends in plant functional traits, equal to 1.0% yr−1, can explain the observed IWUE trends. Thus, at decadal or longer time scales, trait plasticity could potentially influence forest water, carbon, and energy fluxes with profound implications for both the monitoring of temporal changes in plant functional traits and their representation in Earth system models.

Functional trait values, not trait plasticity, drive the invasiveness of Rosa sp. in response to light availability.

Functional trait plasticity in resource capture traits has been suggested as an underlying mechanism promoting invasive species establishment and spread. Earlier studies on this mechanism treat invasiveness as a discrete characteristic (i.e., invasive vs. noninvasive) and do not consider the potential impacts of evolutionary history. In the present study, we used a continuous measure of invasiveness and a phylogenetic framework to quantify the relationship between functional trait expression, plasticity, and invasiveness in Rosa. In a manipulative greenhouse experiment, we evaluated how light availability affects functional traits and their plasticity in Rosa sp. and the out-group species, Potentilla recta, which vary in their invasiveness. Across functional traits, we found no significant relationship between plasticity and invasiveness. However, more invasive roses demonstrated an ability to produce a more branched plant architecture, promoting optimal light capture. Invasiveness also was linked with lower photosynthetic and stomatal conductance rates, leading to increased water-use efficiency (WUE) in more invasive roses. Our results suggest that functional trait values, rather than plasticity, promote invasive rose success, counter to earlier predictions about the role of plasticity in invasiveness. Furthermore, our study indicates that invasive roses demonstrate key functional traits, such as increased WUE, to promote their success in the high-light, edge habitats they commonly invade.

Molecular diversity patterns among various phytoplankton size-fractions in West Greenland in late summer

Arctic regions have experienced pronounced biological and biophysical transformations as a result of global change processes over the last several decades. Current hypotheses propose an elevated impact of those environmental changes on the biodiversity, community composition and metabolic processes of species. The effects on ecosystem function and services, particularly when invasive or toxigenic harmful species become dominant, can be expressed over a wide range of temporal and spatial scales in plankton communities. Our study focused on the comparison of molecular biodiversity of three size-fractions (micro-, nano-, picoplankton) in the coastal pelagic zone of West Greenland and their association with environmental parameters. Molecular diversity was assessed via parallel amplicon sequencing the 28S rRNA hypervariable D1/D2 region. We showed that biodiversity distribution within the area of Uummannaq Fjord, Vaigat Strait and Disko Bay differed markedly within and among size-fractions. In general, we observed a higher diversity within the picoplankton size fraction compared to the nano- and microplankton. In multidimensional scaling analysis, community composition of all three size fractions correlated with cell size, silicate and phosphate, chlorophyll a (chl a) and dinophysistoxin (DTX). Individually, each size fraction community composition also correlated with other different environmental parameters, i.e. temperature and nitrate. We observed a more homogeneous community of the picoplankton across all stations compared to the larger size classes, despite different prevailing environmental conditions of the sampling areas. This suggests that habitat niche occupation for larger-celled species may lead to higher functional trait plasticity expressed as an enhanced range of phenotypes, whereas smaller organisms may compensate for lower potential plasticity with higher diversity. The presence of recently identified toxigenic harmful algal bloom (HAB) species (such as Alexandrium fundyense and A. ostenfeldii) in the area points out the potential risk for this vulnerable ecosystem in a changing world.

Phenotypic plasticity and specialization in clonal versus non-clonal plants: A data synthesis

Reproductive strategies can be associated with ecological specialization and generalization. Clonal plants produce lineages adapted to the maternal habitat that can lead to specialization. However, clonal plants frequently display high phenotypic plasticity (e.g. clonal foraging for resources), factors linked to ecological generalization. Alternately, sexual reproduction can be associated with generalization via increasing genetic variation or specialization through rapid adaptive evolution. Moreover, specializing to high or low quality habitats can determine how phenotypic plasticity is expressed in plants. The specialization hypothesis predicts that specialization to good environments results in high performance trait plasticity and specialization to bad environments results in low performance trait plasticity. The interplay between reproductive strategies, phenotypic plasticity, and ecological specialization is important for understanding how plants adapt to variable environments. However, we currently have a poor understanding of these relationships. In this study, we addressed following questions: 1) Is there a relationship between phenotypic plasticity, specialization, and reproductive strategies in plants? 2) Do good habitat specialists express greater performance trait plasticity than bad habitat specialists? We searched the literature for studies examining plasticity for performance traits and functional traits in clonal and non-clonal plant species from different habitat types. We found that non-clonal (obligate sexual) plants expressed greater performance trait plasticity and functional trait plasticity than clonal plants. That is, non-clonal plants exhibited a specialist strategy where they perform well only in a limited range of habitats. Clonal plants expressed less performance loss across habitats and a more generalist strategy. In addition, specialization to good habitats did not result in greater performance trait plasticity. This result was contrary to the predictions of the specialization hypothesis. Overall, reproductive strategies are associated with ecological specialization or generalization through phenotypic plasticity. While specialization is common in plant populations, the evolution of specialization does not control the nature of phenotypic plasticity as predicted under the specialization hypothesis.

No evidence for larger leaf trait plasticity in ecological generalists compared to specialists

AbstractAimPhenotypic plasticity is hypothesized to contribute to a species’ capacity to occupy broader ranges of conditions and to optimally exploit resource‐rich environments. Although this is supported by case studies of individual species, we do not know whether larger plasticity in functional traits is generally associated with ecological characteristics of species such as their niche breadth or niche position. Here, we test whether there is such a relationship for plasticity in leaf functional traits.LocationCentral Europe.MethodsWe surveyed 110–132 grassland plant species for plasticity in five leaf traits [leaf thickness, leaf greenness, specific leaf area, leaf dry matter content (LDMC) and plant height] and for biomass changes in response to experimental fertilization, shading and waterlogging. Trait plasticity and changes in biomass were compared with species niche characteristics along three environmental axes (light, nutrient and soil moisture) derived from a vegetation‐plot database.ResultsAlthough response of several traits to experimental treatments correlated with niche position and breadth (change in leaf thickness, greenness and biomass in response to fertilization; change in LDMC due to shading; and change in plant height and biomass due to waterlogging), we did not find evidence that species with broader niches or species from resource‐rich environments are more plastic. Ecological generalists even turned out to be less plastic in some traits, including leaf thickness after fertilization and waterlogging. Generalists also displayed smaller plastic response averaged across all five traits (‘composite plasticity’), though the relationship was not statistically significant. This composite plasticity was positively related to absolute change in biomass in all experimental treatments.Main conclusionsOur results suggest that larger species‐level plasticity in leaf traits is not necessarily associated with a capacity to occupy a broader range of environments or with growth in resource‐rich habitats; rather, it may indicate species’ sensitivity to environmental changes.

Biomass Allocation of Stoloniferous and Rhizomatous Plant in Response to Resource Availability: A Phylogenetic Meta-Analysis.

Resource allocation to different functions is central in life-history theory. Plasticity of functional traits allows clonal plants to regulate their resource allocation to meet changing environments. In this study, biomass allocation traits of clonal plants were categorized into absolute biomass for vegetative growth vs. for reproduction, and their relative ratios based on a data set including 115 species and derived from 139 published literatures. We examined general pattern of biomass allocation of clonal plants in response to availabilities of resource (e.g., light, nutrients, and water) using phylogenetic meta-analysis. We also tested whether the pattern differed among clonal organ types (stolon vs. rhizome). Overall, we found that stoloniferous plants were more sensitive to light intensity than rhizomatous plants, preferentially allocating biomass to vegetative growth, aboveground part and clonal reproduction under shaded conditions. Under nutrient- and water-poor condition, rhizomatous plants were constrained more by ontogeny than by resource availability, preferentially allocating biomass to belowground part. Biomass allocation between belowground and aboveground part of clonal plants generally supported the optimal allocation theory. No general pattern of trade-off was found between growth and reproduction, and neither between sexual and clonal reproduction. Using phylogenetic meta-analysis can avoid possible confounding effects of phylogeny on the results. Our results shown the optimal allocation theory explained a general trend, which the clonal plants are able to plastically regulate their biomass allocation, to cope with changing resource availability, at least in stoloniferous and rhizomatous plants.

When variability outperforms the mean: trait plasticity predicts plant cover and biomass in an alpine wetland

Determining which strategies confer a competitive advantage in variable environments with strong short-term resource fluctuations (i.e., seasonal drought and flooding) is critical for understanding ecosystem structure and function, and predicting how ecosystems will respond to global change. Here, we ask how functional traits of plant species vary in response to drought and flooding and whether trait plasticity is a better predictor of plants’ cover than average functional trait values. We transplanted vegetative recruits (similar size and age) of eight common species from a moderately grazed alpine wetland in the Eastern Tibetan Plateau to a common garden where we evaluated the trait responses of these species to drought and flooding. We also measured the coverage, biomass, and height of these plant species in the field communities. The leaf morphological and physiological traits of these eight species generally showed different responses to drought and flood conditions. Variation (plasticity) of leaf morphological or physiological traits rather than average values of these traits were better predictors of cover and biomass in the field community. Species with lower plasticity in leaf morphological traits had higher plasticity in leaf physiology and the higher coverage and biomass in the field community. Our results suggest that the ability to quickly adjust leaf physiology to changing environmental conditions provides an advantage in this ecosystem, where intra-annual rainfall variability is high and thus resource fluctuations are pulsed in time. In contrast, greater functional trait plasticity in leaf morphology was a disadvantage, possibly because the benefits of these adjustments did not outweigh their initial costs.

Plasticity in plant functional traits is shaped by variability in neighbourhood species composition.

Plant functional traits can vary widely as a result of phenotypic plasticity to abiotic conditions. Trait variation may also reflect responses to the identity of neighbours, although not all species are equally responsive to their biotic surroundings. We hypothesized that responses to neighbours are shaped by spatial community patterns and resulting variability in neighbour composition. More precisely, we tested the theoretical prediction that plasticity is most likely to evolve if alternative environments (in this case, different neighbour species) are common and encountered at similar frequencies. We estimated the frequencies of encountering different neighbour species in the field for 27 grassland species and measured the aboveground morphological responses of each species to conspecific vs heterospecific neighbours in a common garden. Responses to neighbour identity were dependent on how frequently the experimental neighbours were encountered by the focal species in their home community, with the greatest plasticity observed in species that encountered both neighbours (conspecific and heterospecific) with high and even frequency. Biotic interactions with neighbouring species can impose selection on plasticity in functional traits, which may feed back through trait divergence and niche differentiation to influence species coexistence and community structure.

Apparent plasticity in functional traits determining competitive ability and spatial distribution: a case from desert.

Species competitive abilities and their distributions are closely related to functional traits such as biomass allocation patterns. When we consider how nutrient supply affects competitive abilities, quantifying the apparent and true plasticity in functional traits is important because the allometric relationships among traits are universal in plants. We propose to integrate the notion of allometry and the classical reaction norm into a composite theoretical framework that quantifies the apparent and true plasticity. Combining the framework with a meta-analysis, a series of field surveys and a competition experiment, we aimed to determine the causes of the dune/interdune distribution patterns of two Haloxylon species in the Gurbantonggut Desert. We found that (1) the biomass allocation patterns of both Haloxylon species in responses to environmental conditions were apparent rather than true plasticity and (2) the allometric allocation patterns affected the plants’ competition for soil nutrient supply. A key implication of our results is that the apparent plasticity in functional traits of plants determines their response to environmental change. Without identifying the apparent and true plasticity, we would substantially overestimate the magnitude, duration and even the direction of plant responses in functional traits to climate change.

Plasticity of functional traits of forb species in response to biodiversity

In spite of increasing recognition that intraspecific variation may play an important role for niche differentiation, which is regarded as a promoter of species coexistence, the extent and structure of functional trait variation in response to plant neighbor diversity is poorly understood. We studied the plasticity of functional traits in vegetative and reproductive shoots of 27 non-legume forb species with different growth forms (reptant, rosulate, semirosulate) in experimental grasslands (Jena Experiment) of varying species richness and functional group composition (with and without legumes). Traits related to whole-shoot structure differed strongly among forb species with different growth forms, while leaf traits associated with light acquisition (specific leaf area, foliar δ13C values) and traits associated with nitrogen nutrition (shoot biomass:N ratios, leaf nitrogen concentrations, foliar δ15N values) were highly plastic within forb species. Plant height generally increased with increasing species richness. Plasticities to increased species richness in leaf traits (leaf length, SLA, foliar δ13C) varied among growth forms and depended on developmental stage. The presence of legumes generally increased plastic responses in light-acquisition traits in the same direction as increasing species richness. Greater tissue nitrogen concentrations and unchanged foliar δ15N values of forb species in the presence of legumes suggested that the fertilizing effect of nitrogen-fixing legumes was due to the supply of unconsumed mineral nitrogen. Stronger correlations between trait means and trait plasticities in size-related traits suggested a functional convergence in response to light competition. A more variable spectrum in the plasticities of traits not associated with plant size indicated a greater functional separation among species. Our results suggest that both interspecific differences and intraspecific trait plasticity affect niche partitioning among forb species and are important for their coexistence in multi-species assemblages.

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 δ(13)C and δ(18)O) 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.