Intraspecific variation in seedling growth responses of a relict tree species Euptelea pleiospermum to precipitation manipulation along an elevation gradient

Improving the accuracy of predictions regarding how plants respond to climate change is crucial to protecting biodiversity. However, little is known about the effects of seed source and elevation on the response of mountain plant species to reductions in precipitation. Here, we collected seeds of a tree species (Euptelea pleiospermum) from three seed sources and carried out a two-growing-season reciprocal transplant experiment with precipitation manipulation at three sites along an elevation gradient in the Shennongjia Mountains, central China. Variations in whole-plant traits, leaf traits, and root traits were investigated. We found that most plant traits of E. pleiospermum seedlings were affected by reductions in precipitation, and responses varied among different elevations and seed sources. Whole-plant traits, root biomass, and leaf traits related to photosynthesis capacity decreased under reduced precipitation treatments at mid and high elevation sites. Thus, climate change induced drought will likely have a negative influence on seedling growth at mid and high elevation regions. In addition, a home-site advantage in whole-plant traits and root traits was observed. However, the responses of leaf traits in most cases were not affected by seed source because of higher phenotypic plasticity. Our results suggested that both local adaptation and phenotypic plasticity were important in seedling growth responses to reduced precipitation. We also highlight the importance of taking intraspecific variation into account when studying the response of plants to changes in climate.

     

Intraspecific phenotypic variability of plant functional traits in contrasting mountain grasslands habitats

Empirical studies that link plants intraspecific variation to environmental conditions are almost lacking, despite their relevance in understanding mechanisms of plant adaptation, in predicting the outcome of environmental change and in conservation. Here, we investigate intraspecific trait variation of four grassland species along with abiotic environmental variation at high spatial resolution (n = 30 samples per species trait and environmental factor per site) in two contrasting grassland habitats in Central Apennines (Italy). We test for phenotypic adaptation between habitats, intraspecific trait-environment relationships within habitats, and the extent of trait and environmental variation. We considered whole plant, clonal, leaf, and seed traits. Differences between habitats were tested using ANOVA and ANCOVA. Trait-environment relationships were assessed using multiple regression models and hierarchical variance partitioning. The extent of variation was calculated using the coefficient of variation. Significant intraspecific differences in trait attributes between the contrasting habitats indicate phenotypic adaptation to in situ environmental conditions. Within habitats, light, soil temperature, and the availability of nitrate, ammonium, magnesium and potassium were the most important factors driving intraspecific trait-environment relationships. Leaf traits and height growth show lower variability than environment being probably more regulated by plants than clonal traits which show much higher variability. We show the adaptive significance of key plant traits leading to intraspecific adaptation of strategies providing insights for conservation of extant grassland communities. We argue that protecting habitats with considerable medium- and small-scale environmental heterogeneity is important to maintain large intraspecific variability within local populations that finally can buffer against uncertainty of future climate and land use scenarios.     

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.     

A Multidimensional Functional Trait Approach Reveals the Imprint of Environmental Stress in Mediterranean Woody Communities

Water availability is one of the most important factors determining species distribution, plant community structure and ecosystem functioning. We explore how the functional structure of Mediterranean woody plant communities varies along a regional gradient of aridity in the Andalusian region (south Spain). We question whether communities located in more arid sites show more similarity in their functional structure when compared with communities located in wetter sites or whether, instead, there is divergence in their functional spaces. We selected five aridity zones (three sampling sites per zone) and measured 13 traits of different functional dimensions (including leaf, stem and root traits) in 74 woody plant species. We quantified functional space differences using the n-dimensional niche space approach (hypervolume). We found a larger functional space for the wetter communities compared with the more arid communities, which showed greater overlap of the trait space occupation. Our results indicate that aridity acts as a key abiotic filter affecting various metrics of the community trait structure, in accordance with the plant economics spectrum. We have also documented consistent variation in the functional space, supporting lower functional diversity under more harsh climatic conditions. The trend of functional space variation along the aridity gradient was different when considering traits from only one plant organ. Thus, the filtering process driving the functional structure of the communities studied here largely depends on the trait axis considered; for example, the root dimension showed considerable variation in wet environments, whereas the leaf dimension exhibited a larger functional space in the drier habitats.     

Genetically based latitudinal variation in Artemisia californica secondary chemistry

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

Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient

I applied a comparative approach to reveal correlated patterns of variation in phenology and seed production in four populations of two annual grasses Hordeum spontaneum and Avena sterilis, sampled in the same environments distributed along an aridity gradient in Israel. The steep aridity gradient in Israel represents two parallel clines of environmental productivity (annual rainfall) and predictability (variation in amount and timing of annual rainfall) that is likely to induce similar responses in natural plant populations distributed along the gradient, if (1) selection is strong, (2) species share the same ecological niche, and (3) there is genetic variation for ecologically important traits. I found in plants of both species (1) ultimate advance in onset of flowering, and (2) more but smaller seeds, with increasing aridity. The broad sense heritabilities of onset of flowering, seed size and seed yield in both species were very high, moderate and low, respectively. It appears that the observed adaptive complex of traits have evolved in both species in response to this specific array of environments.     

Seed mass and dormancy of annual plant populations and communities decreases with aridity and rainfall predictability

Several theoretical and empirical studies have examined the influence of environmental conditions on seed traits and germination strategies of annual species. A positive relationship between seed mass and dormancy has been described for annuals occupying climatically unpredictable ecosystems. Larger-seeded species tend to have higher seedling survival rates, while dormancy allows a bet-hedging strategy in unpredictable environments. Until now, these ideas have been addressed primarily for only one or a few focal species, without considering differences among populations and communities. The novelty of the present study lies in the population and community-level approach, where a comprehensive seed trait database including 158 annual species occurring along a gradient of rainfall variability and aridity in Israel was used to ask the following question: Does average seed mass and dormancy of annual populations and communities decrease with increasing aridity and rainfall unpredictability?Soil seed bank samples were collected at the end of the summer drought, before the onset of the rains, from four plant communities. Germination was tested under irrigated conditions during three consecutive germination seasons to determine the overall seed germinability in each soil sample. Seed mass was obtained from newly produced seeds collected at the study sites in late spring. The community level results showed that, in contrast to common theoretical knowledge, seed mass and dormancy of the dominant annual species decreased with increasing aridity and rainfall variability. Accordingly, a negative correlation was found between seed mass and seed germination fractions. The present study demonstrates that an analysis of seed traits along climatic gradients is significantly improved by approaches that target both population and community levels simultaneously. A critical evaluation sheds new light upon the selective pressures that act on seed ecology of annuals along a climatic gradient and facilitates formulation of more mechanistic hypotheses about factors governing critical seed traits.     

Summer aridity rather than management shapes fitness‐related functional traits of the threatened mountain plant Arnica montana

Semi‐natural mountain grasslands are increasingly exposed to environmental stress under climate change. However, which are the environmental factors that limit plants in these grasslands? Also, is the present management effective against these changes? Fitness‐related functional traits may offer a way to detect changes in performance and provide new insights into their vulnerability to climate change. We investigated changes in performance and variability of functional traits of the mountain grassland target species Arnica montana along a climate gradient in Central German low mountain ranges. This gradient represents at its lower end climate conditions that are expected at its upper end under future climate change. We measured vegetative, generative, and physiological traits to account for multiple ways of plant responses to the environment. Using mixed effects and multivariate models, we evaluated changes in trait values among individuals as well as the variability of their populations in order to assess performance under changing summer aridity and different management regimes. Fitness‐related performance of most traits showed strongly positive associations with reduced summer aridity at higher elevations, while only specific leaf area and leaf dry matter content showed no association. This suggests a higher performance level at less arid montane sites and that the physiological traits are less sensitive to this climate change factor. The coefficient of variation of almost all traits declined steadily with decreasing site aridity. We suggest that this reduced variability indicates a lower environmental stress level for A. montana toward its environmental optimum at montane elevations, especially because the trait performance increased simultaneously. Surprisingly, management factors and habitat characteristics had only low influence on both trait performance and variability. In summary, summer aridity had a stronger effect to shape the trait performance and variability of A. montana under increased environmental stress than management and other habitat characteristics.     

Variation in species‐level plant functional traits over wetland indicator status categories

Wetland indicator status (WIS ) describes the habitat affinity of plant species and is used in wetland delineations and resource inventories. Understanding how species‐level functional traits vary across WIS categories may improve designations, elucidate mechanisms of adaptation, and explain habitat optima and niche. We investigated differences in species‐level traits of riparian flora across WIS categories, extending their application to indicate hydrologic habitat. We measured or compiled data on specific leaf area (SLA ), stem specific gravity (SSG ), seed mass, and mature height of 110 plant species that occur along the Colorado River in Grand Canyon, Arizona. Additionally, we measured leaf δ¹³C, δ¹⁵N, % carbon, % nitrogen, and C/N ratio of 56 species with C3 photosynthesis. We asked the following: (i) How do species‐level traits vary over WIS categories? (ii) Does the pattern differ between herbaceous and woody species? (iii) How well do multivariate traits define WIS categories? (iv) Which traits are correlated? The largest trait differences among WIS categories for herbaceous species occurred for SSG , seed mass, % leaf carbon and height, and for woody species occurred for height, SSG , and δ¹³C. SSG increased and height decreased with habitat aridity for both woody and herbaceous species. The δ¹³C and hence water use efficiency of woody species increased with habitat aridity. Water use efficiency of herbaceous species increased with habitat aridity via greater occurrence of C4 grasses. Multivariate trait assemblages differed among WIS categories. Over all species, SLA was correlated with height, δ¹³C, % leaf N, and C/N; height was correlated with SSG and % leaf C; SSG was correlated with % leaf C. Adaptations of both herbaceous and woody riparian species to wet, frequently inundated habitats include low‐density stem tissue. Adaptations to drier habitats in the riparian zone include short, high‐density cavitation‐resistant stem tissue, and high water use efficiency. The results enhance understanding about using traits to describe plant habitat in riparian systems.     

Morphological responses of Leymus chinensis (Poaceae) to the large-scale climatic gradient along the North-east China Transect (NECT)

A field experiment was conducted to examine the morphological variations of Leymus chinensis along the climatic gradient of the North‐east China Transect (NECT), from 115° to 125° E, in the vicinity of 43.5° N, in north‐eastern China. Ten sites selected for plant sampling along the gradient have approximately uniform theoretical radiation, but differ in precipitation and geographical elevation. The results of analyses showed that vegetative and reproductive shoot heights, flag leaf lengths and widths and seed numbers per inflorescence increased from the west to the east with precipitation, but decreased with aridity. Leaf lengths and widths for most leaf types from 115° to 124° E exhibited little variations. Significant correlations of plant heights, flag leaf lengths and widths and seed numbers per inflorescence with large‐scale climatic variables (e.g. annual precipitation, aridity) and geographical variation (longitude) found in this study indicated that climatic factors have significant effects on some morphological traits of L. chinensis along the NECT.     

Evolvability of between‐year seed dormancy in populations along an aridity gradient

Under global climate change, adaptation to new conditions is crucial for plant species persistence. This requires the ability to evolve in traits that are correlated with changing climatic variables. We studied between‐year seed dormancy, which correlates with environmental variability, and tested for clinal trends in its evolvability along an aridity gradient in Israel. We conducted a germination experiment under five irrigation levels with two dryland winter annuals (Biscutella didyma, Bromus fasciculatus) from four sites along the gradient. Species differed in means and evolvability of dormancy. Biscutella had high dormancy, which significantly increased with aridity but decreased with higher irrigation. In Bromus, dormancy was low, similar among populations, and only marginally affected by irrigation. Evolvability in Biscutella was high and varied among populations, without a clinal trend along the gradient. Conversely, in Bromus, trait evolvability was low and declined with increasing aridity. We argue that changes in evolvability along climatic gradients depend on the relative intensity of stabilizing selection. This may be high in Bromus and not only depends on environmental stress, but also on variability. Our findings point to the importance of measuring evolvability of climate‐related traits across different natural and artificial environments and for many coexisting species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 924–934.     

Patterns of intraspecific trait variation along an aridity gradient suggest both drought escape and drought tolerance strategies in an invasive herb

Background and AimsIn water-limited landscapes, some plants build structures that enable them to survive with minimal water (drought resistance). Instead of making structures that allow survival through times of water limitation, annual plants may invoke a drought escape strategy where they complete growth and reproduction when water is available. Drought escape and resistance each require a unique combination of traits and therefore plants are likely to have a suite of trait values that are consistent with a single drought response strategy. In environments where conditions are variable, plants may additionally evolve phenotypically plastic trait responses to water availability. Invasive annual species commonly occur in arid and semi-arid environments and many will be subject to reduced water availability associated with climate change. Assessing intraspecific trait variation across environmental gradients is a valuable tool for understanding how invasive plants establish and persist in arid environments.MethodsIn this study, we used a common garden experiment with two levels of water availability to determine how traits related to carbon assimilation, water use, biomass allocation and flowering phenology vary in California wild radish populations across an aridity gradient.Key ResultsWe found that populations from arid environments have rapid flowering and increased allocation to root biomass, traits associated with both drought escape and tolerance. Early flowering was associated with higher leaf nitrogen concentration and lower leaf mass per area, traits associated with high resource acquisition. While trait values varied across low- and high-water treatments, these shifts were consistent across populations, indicating no differential plasticity across the aridity gradient.ConclusionsWhile previous studies have suggested that drought escape and drought resistance are mutually exclusive drought response strategies, our findings suggest that invasive annuals may employ both strategies to succeed in novel semi-arid environments. As many regions are expected to become more arid in the future, investigations of intraspecific trait variation within low water environments help to inform our understanding of potential evolutionary responses to increased aridity in invasive species.     

Phenology and polyploidy in annual Brachypodium species (Poaceae) along the aridity gradient in Israel

Local adaptation of plants along environmental gradients provides strong evidence for clinal evolution mediated by natural selection. Plants have developed diverse strategies to mitigate stress, for example, drought escape is a phenological strategy to avoid drought stress, while polyploidy was proposed as a genomic adaptation to stress. Polyploidy as an adaptation to aridity (an environmental parameter integrating temperature and precipitation) was previously documented in annual Brachypodium spp. (Poaceae) in the Western Mediterranean. Here, we examined whether polyploidy or phenology are associated with aridity in annual Brachypodium spp. along the aridity gradient in the Eastern Mediterranean. Using flow cytometry, we determined ploidy levels of plants from natural populations along the Israeli gradient, spanning ∼424 km from mesic Mediterranean to extreme desert climates. In a common garden we recorded time of seedling emergence, flowering and senescence. We tested whether the proportion of allotetraploids in the populations and phenological traits were associated with aridity. Contrary to a previous study in the Western Mediterranean, we found no effect of aridity on the proportion of allotetraploids and diploids within populations. Interestingly, phenology was associated with aridity: time of emergence was later, while flowering and senescence were earlier in desert plants. Our results indicate that in the Eastern Mediterranean, adaptation of Brachypodium to aridity is mediated mainly by phenology, rather than ploidy level. Therefore, we suggest that genome duplication is not the main driver of adaptation to environmental stress; rather, phenological change as a drought escape mechanism may be the major adaptation.     

Invasive species and climate change: <Emphasis Type="Italic">Conyza canadensis</Emphasis> (L.) Cronquist as a tool for assessing the invasibility of natural plant communities along an aridity gradient

The predicted reduction in precipitation in the eastern Mediterranean due to climate change may expose the natural plant communities to invasive species. We assessed whether natural plant communities along an aridity gradient in Israel were resistant to invasion by considering differences in abiotic conditions and community characteristics in these regions. We considered Conyza canadensis as a model plant as it is a common invader in the region. We examined the mechanisms and functional traits of both the plant communities and C. canadensis that promote or discourage invasion. Study sites represented a rainfall gradient with four ecosystem types: mesic Mediterranean, Mediterranean, semiarid and arid. Our results showed that the mechanisms of community invasion resistance varied along the aridity gradient. At the arid and semiarid sites, water deficiency impaired the establishment of C. canadensis. At the mesic Mediterranean site, plant competition had a negative effect on C. canadensis performance, thus greatly reducing the likelihood of its establishment. We conclude that a decrease in regional precipitation due to climate change may not affect intrinsic resistance characteristics of natural plant communities to invasion in the area.     

Trait differences between grass species along a climatic gradient in South and North America

Question: Are trait differences between grasses along a gradient related to climatic variables and/or photosynthetic pathway? Location: Temperate grassland areas of South and North America. Methods: In a common garden experiment, we cultivated C₃ and C₄ grasses from grasslands under different climatic conditions, and we measured a set of 12 plant traits related to size and resource capture and utilization. We described (1) interspecific plant trait differences along a climatic gradient defined by the precipitation and temperature at the location where each species is dominant and (2) the association between those plant trait differences and the photosynthetic pathway of the species. Results: Trait differences between grasses were related to the precipitation at the area where each species is dominant, and to the photosynthetic pathway of the species. Leaf length, leaf width, plant height, leaf area per tiller, specific leaf area, leaf δ¹³C ratio, and nitrogen resorption efficiency increased while leaf dry matter content and nitrogen concentration in senesced leaves decreased as precipitation increased. A proportion of these changes along the gradient was related to the photosynthetic pathway because dominant grass species in cold areas with low precipitation are mainly C₃ and those from warm and wet areas are C₄. Conclusions: A previous worldwide analysis showed that traits of graminoid species measured in situ changed slightly along climatic gradients (< 10% variance explained). In contrast, under a common environment we observed that (1) grass traits changed strongly along a climatic gradient (30‐85% variance explained) and, (2) a proportion of those changes were related to the association between photosynthetic pathway of the species and precipitation.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.     

Constraints on trait combinations explain climatic drivers of biodiversity: the importance of trait covariance in community assembly

Trade‐offs maintain diversity and structure communities along environmental gradients. Theory indicates that if covariance among functional traits sets a limit on the number of viable trait combinations in a given environment, then communities with strong multidimensional trait constraints should exhibit low species diversity. We tested this prediction in winter annual plant assemblages along an aridity gradient using multilevel structural equation modelling. Univariate and multivariate functional diversity measures were poorly explained by aridity, and were surprisingly poor predictors of community richness. By contrast, the covariance between maximum height and seed mass strengthened along the aridity gradient, and was strongly associated with richness declines. Community richness had a positive effect on local neighbourhood richness, indicating that climate effects on trait covariance indirectly influence diversity at local scales. We present clear empirical evidence that declines in species richness along gradients of environmental stress can be due to increasing constraints on multidimensional phenotypes.     

Contrasting intraspecific foliar trait responses to stressful conditions of two rhizomatous granite outcrop species at different scales in southwestern Australia

Plants respond to changing environmental conditions, and their ability to adjust intra‐specifically to such shifts represents an ecological and evolutionary advantage. We studied seven plant traits for two common, rhizomatous granite outcrop species (the fern Cheilanthes austrotenuifolia, and the herb Stypandra glauca) with seasonal foliage during the cooler, wetter winter months at seven sites across an aridity gradient in southwestern Australia. We investigated trait patterns at regional and habitat scale, by investigating changes in trait values along the aridity gradient, and by comparing two different habitats types (sun‐exposed and sheltered). We expected plants occurring in more arid sites and highly irradiated, shallow‐soil (sun‐exposed) habitats, to exhibit traits indicative of more conservative resource acquisition, retention and use strategies. At the habitat scale, we found support for our prediction, with plants in more stressful, sun‐exposed habitats showing traits’ values associated with more conservative strategies (especially for water), such as smaller plants, denser leaves, higher foliar δ¹³C and C/N. However, at the regional scale many traits displayed the opposite pattern, suggesting less conservative resource acquisition in more arid sites. This evidence was particularly pronounced for specific leaf area (SLA), which exhibited a significant, positive relationship with increasing aridity. We suggest that the unexpected regional trends in foliar traits relate to shorter lived, faster growing leaves linked to highly efficient resource acquisition and use strategies during the shorter growing season in the more arid regions. These highly exploitative strategies may enable plants to avoid climate extremes, that is, hot and dry periods in the more arid sites. Our findings of contrasting foliar traits responses at different scales support the importance of multi‐scale approaches to quantify the role of intraspecific trait variability.     

Shifts in fine root traits within and among species along a fine-scale hydrological gradient

Background and AimsLessons from above-ground trait ecology and resource economics theory may not be directly translatable to below-ground traits due to differences in function, trade-offs and environmental constraints. Here we examine root functional traits within and across species along a fine-scale hydrological gradient. We ask two related questions: (1) What is the relative magnitude of trait variation across the gradient for within- versus among-species variation? (2) Do correlations among below-ground plant traits conform with predictions from resource-economic spectrum theory?MethodsWe sampled four below-ground fine-root traits (specific root length, branching intensity, root tissue density and root dry matter content) and four above-ground traits (specific leaf area, leaf size, plant height and leaf dry matter content) in vascular plants along a fine-scale hydrological gradient within a wet heathland community in south-eastern Australia. Below-ground and above-ground traits were sampled both within and among species.Key ResultsRoot traits shifted both within and among species across the hydrological gradient. Within- and among-species patterns for root tissue density showed similar declines towards the wetter end of the gradient. Other root traits showed a variety of patterns with respect to within- and among-species variation. Filtering of species has a stronger effect compared with the average within-species shift: the slopes of the relationships between soil moisture and traits were steeper across species than slopes of within species. Between species, below-ground traits were only weakly linked to each other and to above-ground traits, but these weak links did in some cases correspond with predictions from economic theory.ConclusionsOne of the challenges of research on root traits has been considerable intraspecific variation. Here we show that part of intraspecific root trait variation is structured by a fine-scale hydrological gradient, and that the variation aligns with among-species trends in some cases. Patterns in root tissue density are especially intriguing and may play an important role in species and individual response to moisture conditions. Given the importance of roots in the uptake of resources, and in carbon and nutrient turnover, it is vital that we establish patterns of root trait variation across environmental gradients.     

Leaf traits and performance vary with plant age and water availability in Artemisia californica

Background and AimsLeaf functional traits are strongly tied to growth strategies and ecological processes across species, but few efforts have linked intraspecific trait variation to performance across ontogenetic and environmental gradients. Plants are believed to shift towards more resource-conservative traits in stressful environments and as they age. However, uncertainty as to how intraspecific trait variation aligns with plant age and performance in the context of environmental variation may limit our ability to use traits to infer ecological processes at larger scales.MethodsWe measured leaf physiological and morphological traits, canopy volume and flowering effort for Artemisia californica (California sagebrush), a dominant shrub species in the coastal sage scrub community, under conditions of 50, 100 and 150 % ambient precipitation for 3 years.Key ResultsPlant age was a stronger driver of variation in traits and performance than water availability. Older plants demonstrated trait values consistent with a more conservative resource-use strategy, and trait values were less sensitive to drought. Several trait correlations were consistent across years and treatments; for example, plants with high photosynthetic rates tended to have high stomatal conductance, leaf nitrogen concentration and light-use efficiency. However, the trade-off between leaf construction and leaf nitrogen evident in older plants was absent for first-year plants. While few traits correlated with plant growth and flowering effort, we observed a positive correlation between leaf mass per area and performance in some groups of older plants.ConclusionsOverall, our results suggest that trait sensitivity to the environment is most visible during earlier stages of development, after which intraspecific trait variation and relationships may stabilize. While plant age plays a major role in intraspecific trait variation and sensitivity (and thus trait-based inferences), the direct influence of environment on growth and fecundity is just as critical to predicting plant performance in a changing environment.