Intraspecific Leaf Trait Research Articles

Intraspecific leaf trait variation mediates edge effects on litter decomposition rate in fragmented forests.

There is strong trait dependence in species-level responses to environmental change and their cascading effects on ecosystem functioning. However, there is little understanding of whether intraspecific trait variation (ITV) can also be an important mechanism mediating environmental effects on ecosystem functioning. This is surprising, given that global change processes such as habitat fragmentation and the creation of forest edges drive strong trait shifts within species. On 20 islands in the Thousand Island Lake, China, we quantified intraspecific leaf trait shifts of a widely distributed shrub species, Vaccinium carlesii, in response to habitat fragmentation. Using a reciprocal transplant decomposition experiment between forest edge and interior on 11 islands with varying areas, we disentangled the relative effects of intraspecific leaf trait variation versus altered environmental conditions on leaf decomposition rates in forest fragments. We found strong intraspecific variation in leaf traits in response to edge effects, with a shift toward recalcitrant leaves with low specific leaf area and high leaf dry matter content from forest interior to the edge. Using structural equation modeling, we showed that such intraspecific leaf trait response to habitat fragmentation had translated into significant plant afterlife effects on leaf decomposition, leading to decreased leaf decomposition rates from the forest interior to the edge. Importantly, the effects of intraspecific leaf trait variation were additive to and stronger than the effects from local environmental changes due to edge effects and habitat loss. Our experiment provides the first quantitative study showing that intraspecific leaf trait response to edge effects is an important driver of the decrease in leaf decomposition rate in fragmented forests. By extending the trait-based response-effect framework toward the individual level, intraspecific variation in leaf economics traits can provide the missing functional link between environmental change and ecological processes. These findings suggest an important area for future research on incorporating ITV to understand and predict changes in ecosystem functioning in the context of global change.

Intraspecific Leaf Trait Variation across and within Five Common Wine Grape Varieties.

Variability in traits forming the Leaf Economics Spectrum (LES) among and within crop species plays a key role in governing agroecosystem processes. However, studies evaluating the extent, causes, and consequences of within-species variation in LES traits for some of the world’s most common crops remain limited. This study quantified variations in nine leaf traits measured across 90 vines of five common wine grape (Vitis vinifera L.) varieties at two growth stages (post-flowering and veraison). Grape traits in these varieties covary along an intraspecific LES, in patterns similar to those documented in wild plants. Across the five varieties evaluated here, high rates of photosynthesis (A) and leaf nitrogen (N) concentrations were coupled with low leaf mass per area (LMA), whereas the opposite suite of traits defined the “resource-conserving end” of this intraspecific LES in grape. Variety identity was the strongest predictor of leaf physiological (A) and morphological traits (i.e., leaf area and leaf mass), whereas leaf chemical traits and LMA were best explained by growth stage. All five varieties expressed greater resource-conserving trait syndromes (i.e., higher LMA, lower N, and lower Amass) later in the growing season. Traits related to leaf hydraulics, including instantaneous water-use efficiency (WUE), were unrelated to LES and other resource capture traits, and were better explained by spatial location. These results highlight the relative contributions of genetic, developmental, and phenotypic factors in structuring trait variation in the five wine grape varieties evaluated here, and point to a key role of domestication in governing trait relationships in the world’s crops.

Life stages and habitat types alter the relationships of tree growth with leaf traits and soils in an old-growth temperate forest

• Growth of saplings is more affected by leaf traits and soil variation than that of adults in broadleaf trees. • Growth of adults is more affected by leaf traits and soil variation than that of saplings in coniferous trees. • Absolute growth rate is more sensitive to leaf traits and soil variation than relative growth rate. • Leaf traits showed different effects on broadleaf sapling growth in different habitat types. • Soil variation had positive effects on broadleaf sapling and coniferous adult growth in the high-nutrient habitat type. Knowledge of the leaf trait–growth rate relationship is essential for assessing plant physiological performance in forest ecosystems. Intraspecific leaf trait variation may have different effects on tree growth at various spatial and temporal scales. Plant functional-group and life-stage approaches have the potential to clarify the specific roles to understand the biodiversity–ecosystem functioning relationship. Our research determined the significance of leaf traits and soil variation in broadleaf and coniferous tree growth across different life stages and habitat types. In a 9 ha Liangshui forest dynamic plot in Northeast China, we used linear mixed-effects models (LMMs) to model tree growth as a function of leaf traits and soil variables in three habitat types with different soil organic matter and slope conditions. Tree growth in the broadleaf saplings was more significantly influenced by leaf traits and soil variation than that in the adults. On the other hand, the growth of coniferous adults was more significantly influenced by leaf traits and soil variation than that of the saplings. The influence of leaf traits on the growth of broadleaf saplings was dependent on the habitat type. Soil variation had positive effects on the growth of broadleaf saplings and coniferous adults in the high-nutrient habitat type. Different functional types can provide different insights into the influence of leaf traits on tree growth. To apply leaf traits to forest management, the relationships of plant functional types, life stages, and local environments with tree growth need to be carefully considered, particularly in mixed-species forests with varying tree sizes.

Differential nutrient limitation and tree height control leaf physiology, supporting niche partitioning in tropical dipterocarp forests

Abstract Revealing the mechanisms of environmental niche partitioning within lowland tropical forests is important for understanding the drivers of current species distributions and potential vulnerability to environmental change. Tropical forest structure and species composition change across edaphic gradients in Borneo over short distances. However, our understanding of how edaphic conditions affect tree physiology and whether these relationships drive niche partitioning within Bornean forests remains incomplete. This study evaluated how leaf physiological function changes with nutrient availability across a fine‐scale edaphic gradient and whether these relationships vary according to tree height. Furthermore, we tested whether intraspecific leaf trait variation allows generalist species to populate a wider range of environments. We measured leaf traits of 218 trees ranging in height from 4 to 66 m from 13 dipterocarp species within four tropical forest types (alluvial, mudstone, sandstone and kerangas) occurring along an <5 km edaphic gradient in North Borneo. The traits measured included saturating photosynthesis (Asat), maximum photosynthetic capacity (Vcmax), leaf dark respiration (Rleaf), leaf mass per area (LMA), leaf thickness, minimum stomatal conductance (gdark) and leaf nutrient concentrations (N, P, Ca, K and Mg). Across all species, leaf traits varied consistently in response to soil nutrient availability across forest types except Rleaf_mass, [Mg]leaf and [Ca]leaf. Changes in photosynthesis and respiration rates were related to different leaf nutrients across forest types, with greater nutrient‐use efficiency in more nutrient‐poor environments. Generalist species partially or fully compensated reductions in mass‐based photosynthesis through increasing LMA in more nutrient‐poor environments. Leaf traits also varied with tree height, except Vcmax_mass, but only in response to height‐related modifications of leaf morphology (LMA and leaf thickness). These height–trait relationships did not vary across the edaphic gradient, except for Asat, [N]leaf, [P]leaf and [K]leaf. Our results highlight that modification of leaf physiological function and morphology act as important adaptations for Bornean dipterocarps in response to edaphic and vertical environmental gradients. Meanwhile, multiple nutrients appear to contribute to niche partitioning and could drive species distributions and high biodiversity within Bornean forest landscapes. Read the free Plain Language Summary for this article on the Journal blog.

Environmental variation drives the decoupling of leaf and root traits within species along an elevation gradient.

Plant performance is enhanced by balancing above- and below-ground resource uptake through the intraspecific adjustment of leaf and root traits. It is assumed that these organ adjustments are at least partly coordinated, so that analogous leaf and root traits broadly covary. Understanding the extent of such intraspecific leaf-root trait covariation would strongly contribute to our understanding of how plants match above- and below-ground resource use strategies as their environment changes, but comprehensive studies are lacking. We measured analogous leaf and root traits from 11 species, as well as climate, soil and vegetation properties along a 1000-m elevation gradient in the French Alps. We determined how traits varied along the gradient, to what extent this variation was determined by the way different traits respond to environmental cues acting at different spatial scales (i.e. within and between elevations), and whether trait pairs covaried within species. Leaf and root trait patterns strongly diverged: across the 11 species along the gradient, intraspecific leaf trait patterns were largely consistent, whereas root trait patterns were highly idiosyncratic. We also observed that, when compared with leaves, intraspecific variation was greater in root traits, due to the strong effects of the local environment (i.e. at the same elevation), while landscape-level effects (i.e. at different elevations) were minor. Overall, intraspecific trait correlations between analogous leaf and root traits were nearly absent. Our study suggests that environmental gradients at the landscape level, as well as local heterogeneity in soil properties, are the drivers of a strong decoupling between analogous leaf and root traits within species. This decoupling of plant resource acquisition strategies highlights how plants can exhibit diverse whole-plant acclimation strategies to modify above- and below-ground resource uptake, improving their resilience to environmental change.

Leaf hydraulics coordinated with leaf economics and leaf size in mangrove species along a salinity gradient

Independence among leaf economics, leaf hydraulics and leaf size confers plants great capability in adapting to heterogeneous environments. However, it remains unclear whether the independence of the leaf traits revealed across species still holds within species, especially under stressed conditions. Here, a suite of traits in these dimensions were measured in leaves and roots of a typical mangrove species, Ceriops tagal, which grows in habitats with a similar sunny and hot environment but different soil salinity in southern China. Compared with C. tagal under low soil salinity, C. tagal under high soil salinity had lower photosynthetic capacity, as indicated directly by a lower leaf nitrogen concentration and higher water use efficiency, and indirectly by a higher investment in defense function and thinner palisade tissue; had lower water transport capacity, as evidenced by thinner leaf minor veins and thinner root vessels; and also had much smaller single leaf area. Leaf economics, hydraulics and leaf size of the mangrove species appear to be coordinated as one trait dimension, which likely stemmed from co-variation of soil water and nutrient availability along the salinity gradient. The intraspecific leaf trait relationship under a stressful environment is insightful for our understanding of plant adaption to the multifarious environments.

Size-dependent and environment-mediated shifts in leaf traits of a deciduous tree species in a subtropical forest.

AimsUnderstanding the joint effects of plant development and environment on shifts of intraspecific leaf traits will advance the understandings of the causes of intraspecific trait variation. We address this question by focusing on a widespread species Clausena dunniana in a subtropical broad‐leaved forest.MethodsWe sampled 262 individuals of C. dunniana at two major topographic habitat types, the slope and hilltop, within the karst forests in Maolan Nature Reserve in southwestern China. We measured individual plant level leaf traits (i.e., specific leaf area (SLA), leaf area, leaf dry‐matter content (LDMC), and leaf thickness) that are associated with plant resource‐use strategies. We adopted a linear mixed‐effects model in which the plant size (i.e., the first principal component of plant basal diameter and plant height) and environmental factors (i.e., topographic habitat, canopy height, and rock‐bareness) were used as independent variables, to estimate their influences on the shifts of leaf traits.Key ResultsWe found that (1) plant size and the environmental factors independently drove the intraspecific leaf trait shifts of C. dunniana, of which plant size explained less variances than environmental factors. (2) With increasing plant size, C. dunniana individuals had increasingly smaller SLA but larger sized leaves. (3) The most influential environmental factor was topographic habitat; it drove the shifts of all the four traits examined. Clausena dunniana individuals on hilltops had leaf traits representing more conservative resource‐use strategies (e.g., smaller SLA, higher LDMC) than individuals on slopes. On top of that, local‐scale environmental factors further modified leaf trait shifts.ConclusionsPlant size and environment independently shaped the variations in intraspecific leaf traits of C. dunniana in the subtropical karst forest of Maolan. Compared with plant size, the environment played a more critical role in shaping intraspecific leaf trait variations, and potentially also the underlying individual‐level plant resource‐use strategies.

Evaluating different methods for retrieving intraspecific leaf trait variation from hyperspectral leaf reflectance

Leaf mass per area (LMA), leaf dry matter content (LDMC) and leaf water content/ equivalent water thickness (EWT) are commonly used functional plant traits in ecology. Whereas spectroscopy has recently proven to be a powerful tool to collect such functional trait information across large scales, it remains unclear whether these reflectance-based trait predictions are accurate enough to reliably model trait variation at the intraspecific level (i.e. across individuals of one species). We explored the potential of hyperspectral leaf reflectance-based methods to predict LMA, LDMC and EWT at the intraspecific level for two herbs (Hieracium umbellatum and Jacobaea vulgaris) and two shrubs (Rosa rugosa and Rubus caesius), based on 2400 leaf samples. More specifically we tested i) inversion of the PROSPECT-D radiative transfer model, ii) a generic PLSR approach using the multibiome LMA PLSR model and iii) a data-specific PLSR approach at the species level. For the latter approach we furthermore assessed both model transferability across species and the trade-off between sample size and model accuracy. Although the PROSPECT-D model inversion and the multibiome LMA PLSR model were relatively accurate for intraspecific LMA predictions of shrubs (R2 > 71 and 76%, respectively, however NRMSE = 33–47%), their performance was lower for herbs (R2 < 61%, NRMSE = 28–50%). PROSPECT-D was furthermore slightly less successful in retrieving EWT at the intraspecific level (R2 < 70%, NRMSE = 16–43%), and unsuccessful in retrieving LDMC through combining LMA and EWT inversion results (R2 < 10%, NRMSE = 9–192%). The highest correlation accuracy was obtained for all three traits with the species-specific PLSR models (R2 > 70%, NRMSE < 10%). If high predictive accuracy is needed, we thus suggest the use of species-specific PLSR models. The training of species-specific PLSR models comes at the cost of a needed sample size of 100–160 leaves however, depending on the trait. Although transferability of species-specific PLSR models seems limited overall, our results suggest potentially high transferability across herbaceous species.

Intra-specific leaf trait variability of F. sylvatica, Q. petraea and P. abies in response to inter-specific competition and implications for forest functioning.

Variability in functional traits (FT) is increasingly used to understand the mechanisms behind tree species interactions and ecosystem functioning. In order to explore how FT differ due to interactions between tree species and its influence on stand productivity and other ecological processes, we examined the effects of tree species composition on the intra-specific variability of four widely measured FT: specific leaf area, leaf nitrogen content, leaf angle and stomatal conductance response to vapor pressure deficit. This study focused on three major central European tree species: European beech (Fagus sylvatica L.), Sessile oak (Quercus petraea Liebl.) and Norway spruce (Picea abies [L.] H. Karst.). Each species was examined in monoculture and two-species mixtures in the 13-year-old tree biodiversity experiment BIOTREE-Kaltenborn. Trait distributions and linear mixed models were used to analyze the effect of species mixing, tree size and stand variables on the intra-specific FT variability. A significant effect of branch height on most traits and species indicated a vertical gradient of foliar trait frequently related to light availability. Beech and oak showed a high overall trait variability and sensitivity to species mixing and stand basal area, while the trait variability of spruce was limited. Greater shifts in trait distributions due to mixing were found in specific leaf area for oak and leaf nitrogen content for beech. Thus intra-specific variability of key leaf traits was already influenced at this young development stage by inter-specific interactions. Finally, we used the 3-PG (Physiological Processes Predicting Growth) process-based forest growth model to show that the measured intra-specific variability on single FT values could influence stand productivity, light absorption and transpiration, although the net effect depends on the considered trait and the species composition of the mixture. The results of this study will aid better understanding of the effects of inter-specific competition on intra-specific FT variability, which has implications for the parameterization of process-based forest growth models and our understanding of ecosystem functioning.

Intra-specific leaf trait variability controls leaf decomposition of Vitis vinifera L. cultivars in streams

Intra-specific leaf trait variability controls leaf decomposition of Vitis vinifera L. cultivars in streams

Intra-specific leaf trait responses to species richness at two different local scales

Plant functional traits, especially leaf traits, are accepted proxies for ecosystem properties. Typically, they are measured at the species level, neglecting within-species variation. While there is extensive knowledge about functional trait changes (both within and across species) along abiotic gradients, little is known about biotic influences, in particular at local scales. Here, we used a large biodiversity-ecosystem functioning experiment in subtropical China to investigate intra-specific trait changes of 16 tree species as a response to species richness of the local neighbourhood. We hypothesized that because of positive complementarity effects, species shift their leaf traits towards a more acquisitive growth strategy, when species richness of the local neighbourhood is higher. The trait shift should be most pronounced, when a focal tree's closest neighbour is from a different species, but should still be detectable as a response to species richness of the directly surrounding tree community. Consequently, we expected that trees with a con-specific closest neighbour have the strongest response to species richness of the surrounding tree community, i.e., the steepest increase of acquisitive traits. Our results indicate that species diversity promoted reduced competition and complementarity in resource use at both spatial scales considered. In addition, the closest neighbour had considerably stronger effects than the surrounding tree community. As expected, trees with a con-specific nearest neighbour showed the strongest trait shifts. However, the predicted positive effect of local hetero-specificity disappeared at the highest diversity levels of the surrounding tree community, potentially resulting from a higher probability to meet a strong competitor in a diverse environment. Our findings show that leaf traits within the same species vary not only in response to changing abiotic conditions, but also in response to local species richness. This highlights the benefit of including within-species trait variation when analysing relationships between plant functional traits and ecosystem functions.

Variations in leaf morphological and chemical traits in response to life stages, plant functional types, and habitat types in an old-growth temperate forest

Intraspecific leaf trait variations are becoming a topic of interest for many ecologists because individual-based traits are essentially the drivers of variations at the community level. Six coexisting major tree species in an old-growth temperate forest, Northeast China (i.e., Abies nephrolepis, Pinus koraiensis, Acer mono, Fraxinus mandshurica, Tilia amurensis, and Ulmus laciniata) were sampled, and three habitat types (i.e., Hab I: high soil organic carbon with a moderate slope; Hab II: low soil organic carbon with a gentle slope; and Hab III: low soil organic carbon with a strong slope) were used in the plot. We performed a two-way ANOVA to compare the specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen content (LNC), leaf phosphorus content (LPC), and leaf carbon content (LCC) between saplings (1 < DBH ≤ 5 cm) and adults (DBH ≥ 10 cm) and between habitat types within species. We simultaneously evaluated the effects of life stage, plant functional type, and habitat type on the six leaf traits. Our results showed that life stage and habitat type had varied influences on the leaf traits of the six species. Life stage was an important determinant for all leaf traits. Plant functional type was included in the best models for SLA, LNC, and LCC. Habitat type had a greater influence on LDMC than SLA. Meanwhile, habitat type had a greater influence on LNC and LPC than LCC. The correlation between leaf traits with local environmental factors varied across different plant functional types and life stages. We suggest conducting individual-based analyses of leaf trait variations according to plant functional type and life stage to understand the plant life strategies along an environmental gradient may improve understanding of the forest dynamics in an old-growth temperate forest.

Leaf trait variation and decomposition in short-rotation woody biomass crops under agroforestry management

The integration of trees into agricultural landscapes has a strong influence on plant-soil interactions, with major impacts on rates of nutrient cycling across multiple spatial scales. Shifts in abiotic conditions owing to agroforestry practices, such as changes in soil nutrient and microclimatic conditions, are commonly identified as primary mechanisms for these changes. Yet agroforestry systems are also likely to influence litter decomposition rates through their influence on morphological and chemical traits of crop leaves and residues. However, explicit evaluations of how agroforestry management results in intraspecific leaf trait plasticity, and how this variation then links with litter decomposition, remain sparse. Using a 30-year-old agroforestry system intercropped with short-rotation willow (Salix dasyclados), we i) examined the effects of agroforestry management (i.e. N2-fixing trees vs. non N2-fixing trees vs. monoculture) and distance from tree row on key crop (willow) leaf morphological (i.e. leaf area, specific-leaf area [SLA]) and chemical (leaf carbon (C) and nitrogen (N) concentration) traits, and then assessed ii) how this intraspecific leaf trait variability influenced decomposition rates. Willow leaves within agroforestry treatments exhibited significantly greater leaf area (p = 0.01), SLA (p = 0.002), and leaf N (p = 0.001) than willow leaves in monoculture. Following a 56-day incubation, willow leaves from agroforestry treatments did not decompose more quickly than those in the monoculture. Our results indicate high intraspecific variation in willow leaf traits owing to agroforestry management. However, intraspecific variation in willow leaf traits did not correspond to statistically significant differences in decomposition rates. We show that variation in willow traits near tree rows in agroforestry systems translates into limited variability in agroecosystems processes such as litter decomposition (as compared to monoculture arrangements). Our results have implications for optimal nutrient management in agroforestry systems.

Strong intraspecific trait variation in a tropical dominant tree species along an elevational gradient

Functional trait variation of plant species includes both inter- and intraspecific variation; however, trait-based plant ecology generally considers only interspecific variation while ignoring intraspecific variation. One reason for this neglect is that intraspecific variation may be negligible when compared to interspecific variation; however, direct comparisons between inter- and intraspecific variation of plant species are lacking, especially in tropical forests. Here we investigated intraspecific leaf trait variation (leaf area, specific leaf area, leaf thickness, leaf density, leaf chlorophyll content) of Pittosporopsis kerrii Craib (Icacinaceae), the most abundant tree species in the Xishuangbanna tropical seasonal rainforest in southwestern China, along an elevational gradient (703–824 m). We found a substantial range of intraspecific variation in P. kerrii that was never less than 22.1% of range of the interspecific variation among 462 tree species reported before in the same community. Moreover, with increased elevation, both leaf thickness and density increased and specific leaf area decreased significantly. It could be more important for the individuals of P. kerrii to produce thicker and denser leaves to tolerate environmental stress (e.g. soil water availability) rather than having high growth rates at the places with higher elevation in the Xishuangbanna tropical seasonal rainforest.

Does trait variation within broadly distributed species mirror patterns across species? A case study in Puerto Rico.

Although populations are phenotypically diverse, the majority of trait-based studies have focused on examining differences among species. The justification for this broadly applied approach is based on the assumption that differences among species are always greater than within species. This is likely true for local communities, but species are often broadly distributed across a wide range of environments and patterns of intraspecific variation might surpass differences among species. Therefore, an appropriate interpretation of the functional diversity requires an assessment of patterns of trait variation across different ecological scales. In this study, we examine and characterize patterns of leaf trait variation for species that are broadly distributed along an elevational gradient. We focus on seven leaf traits that represent a main axis of functional differentiation in plants reflecting the balance between photosynthetic efficiency, display, and stomatal conductance. We evaluated patterns of trait variance across ecological scales (elevation, species, populations, and individuals) and examined trait covariance at both within species and across species levels, along the elevation gradient. Our results show three key patterns: (1) intraspecific leaf trait variation for broadly distributed species is comparable to the interspecific trait variation, (2) the trait variance structure is highly variable across species, and (3) trait coordination between pairs of leaf traits is evident across species along the gradient, but not always within species. Combined, our results show that trait coordination and covariance are highly idiosyncratic across broadly distributed and co-occurring species, indicating that species may achieve similar functional roles even when exhibiting different phenotypes. This result challenges the traditional paradigm of functional ecology that assumes single trait values as optimal solutions for environments. In conclusion, patterns of trait variation both across and within species should be considered in future studies that assess trade-offs among traits over environmental gradients.

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 (δ13 C). We found LMA, Narea, Nmass and δ13 C 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 δ13 C 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.

The interplay among intraspecific leaf trait variation, niche breadth and species abundance along light and soil nutrient gradients

It is assumed that widespread, generalist species have high phenotypic variation, but we know little about how intraspecific trait variation (ITV) relates to species abundance and niche breadth. In the temperate rainforest of southern Chile, we hypothesized that species with wide niche breadth would exhibit 1) high among‐plot ITV, 2) a strong relationship between trait values and the environment, and 3) a close fit between traits and local environment trait optima. We measured leaf functional traits (leaf area, LMA, leaf N and P concentrations) of saplings in woody species, and compared the relative abundance of each species with its niche breadth, measured as the range of light, soil N and P availability. We used the slope of the linear regression of species’ trait–environment relationships to assess the strength and direction of these relationships, and measured the degree to which species’ trait values track the environmental optimum across plots. In some cases, species having wide niche breadth had high ITV in leaf N and also matched traits (LMA and leaf P) to local optima along the light gradient; they also had high ITV in general and matched leaf P to local optima along the soil P gradient. The relationship between species with wide niche breadth and the strength of intraspecific trait–environment relationships was generally weak and varied depending on the niche dimension and trait in question. Species varied considerably in the strength of trait–environment relationships and total magnitude of ITV, and this variation was not generally strongly related to species abundances or niche breadth patterns. In conclusion, trait variation at the community level is not driven by a few abundant, widely distributed species, but depends on the aggregate trait responses of both abundant and rare species. This makes it difficult to scale individual species trait responses up to the community level.

Intraspecific trait variation and the leaf economics spectrum across resource gradients and levels of organization.

Understanding patterns of functional trait variation across environmental gradients offers an opportunity to increase inference in the mechanistic causes of plant community assembly. The leaf economics spectrum (LES) predicts global tradeoffs in leaf traits and trait-environment relationships, but few studies have examined whether these predictions hold across different levels of organization, particularly within species. Here, we asked (1) whether the main assumptions of the LES (expected trait relationships and shifts in trait values across resource gradients) hold at the intraspecific level, and (2) how within-species trait correlations scale up to interspecific or among-community levels. We worked with leaf traits of saplings of woody species growing across light and soil N and P availability gradients in temperate rainforests of southern Chile. We found that ITV accounted for a large proportion of community-level variation in leaf traits (e.g., LMA and leaf P) and played an important role in driving community-level shifts in leaf traits across environmental gradients. Additionally, intraspecific leaf trait relationships were generally consistent with interspecific and community-level trait relationships and with LES predictions-e.g., a strong negative intraspecific LMA-leaf N correlation-although, most trait relationships varied significantly among species, suggesting idiosyncrasies in the LES at the intraspecific level.

Intraspecific leaf trait variability along a boreal-to-tropical community diversity gradient.

Disentangling the mechanisms that shape community assembly across diversity gradients is a central matter in ecology. While many studies have explored community assembly through species average trait values, there is a growing understanding that intraspecific trait variation (ITV) can also play a critical role in species coexistence. Classic biodiversity theory hypothesizes that higher diversity at species-rich sites can arise from narrower niches relative to species-poor sites, which would be reflected in reduced ITV as species richness increases. To explore how ITV in woody plant communities changes with species richness, we compiled leaf trait data (leaf size and specific leaf area) in a total of 521 woody plant species from 21 forest communities that differed dramatically in species richness, ranging from boreal to tropical rainforests. At each forest, we assessed ITV as an estimate of species niche breadth and we quantified the degree of trait overlap among co-occurring species as a measure of species functional similarity. We found ITV was relatively invariant across the species richness gradient. In addition, we found that species functional similarity increased with diversity. Contrary to the expectation from classic biodiversity theory, our results rather suggest that neutral processes or equalizing mechanisms can be acting as potential drivers shaping community assembly in hyperdiverse forests.

Tree species richness induces strong intraspecific variability of beech (Fagus sylvatica) leaf traits and alleviates edaphic stress

Manipulating stand composition is an important management tool that foresters can use to affect the nature of forests and ecosystem processes. In mixed stands, interspecific interactions among trees can cause changes in tree performances. Nevertheless, these interactions are context dependent (cf. stress-gradient hypothesis, SGH). We thus investigated how intraspecific functional changes in leaf trait (19 traits) of European beech (Fagus sylvatica) were influenced by stand composition. We compared pure beech stands with four mixed stands containing from one to three additional tree species along a gradient of edaphic stress (gradient of soil water-holding capacity and rooting depth). First, we demonstrated that stand composition induced strong intraspecific leaf trait variation in beech for LDMC, LMA, phenolic compounds, leaf pH and magnesium concentration, suggesting higher nutrient acquisition by more diverse stands. Nevertheless, these results were modulated by edaphic stress. Mixed stands only conferred an advantage in relatively stressed sites (luvisol and leptosol). Besides, the addition of oak to beech stands had unexpected negative effects in sites with less severe stress (cambisol) as indicated by the null or positive LogRR of LMA, LDMC and phenolics. This study found that stand composition is an important though often-overlooked driver of intraspecific variability in leaf quality, and potentially reflects changes in beech tree physiology and productivity. Our results also suggest that positive interactions prevail in sites with stressful conditions. Such validation of the SGH is rare in natural or managed mature forests. Lastly, we strongly recommend that forest managers consider stand composition and abiotic factors when implementing forest growth models to improve their yield predictions.