Leaf Economic Traits Research Articles

Strong conservatism in leaf anatomical traits and their multidimensional relationships with leaf economic traits in grasslands under different stressful environments

BackgroundPlant traits and plant adaptive strategies have been affected by the increasing intensity and severity of environmental changes. Given the uncertainty surrounding future environmental conditions, investigating plant trait variations under various stresses is crucial for unraveling plant survival strategies. Leaf anatomical traits are closely responsible for plants’ photosynthesis, respiration and transpiration. However, knowledge of how the multi-species leaf anatomical traits varied in extremely and moderately stressful environments is limited. Our objective was to compare the variation of leaf anatomic traits and adaptation strategies in two different stressful regions of the Qinghai-Tibet Plateau (TP) and Mongolian Plateau (MP) of China.MethodsWe sampled ten sites in each of the two regions (MP and TP) along an environmental gradient. Seven leaf anatomical traits and two leaf economic traits were measured for all leaf samples. Leaf anatomical traits include the traits related to leaf physiological processes (mesophyll thickness (MT), palisade tissue thickness (PT), spongy tissue thickness (ST), palisade-spongy tissue thickness ratio (PST) and epidermal thickness (ET)) and the traits related to trait construction investment (epiderm-leaf thickness ratio (ET/LT) and mesophyll-leaf thickness ratio (MT/LT)). Leaf economic traits include specific leaf area (SLA) and leaf nitrogen content (LN).ResultsThe results revealed that leaf anatomical traits in the TP exhibited greater phylogenetic conservation with thicker structures, being less susceptible to environmental impacts than those in the MP. Additionally, the leaf anatomical and economic traits decoupled both in the MP and TP.ConclusionThese findings highlight that plants adopt diverse strategies to cope with extremely and moderately environmental stresses, but multidimensional trait patterns are generally favored in stressful environments.

Linkage between leaf anatomical structure and key leaf economic traits across co-existing species in temperate forests

Linkage between leaf anatomical structure and key leaf economic traits across co-existing species in temperate forests

Coordination of water use strategies and leaf economic traits in coexisting exotic and native woody species from evergreen and deciduous broadleaf forests

The leaf economics spectrum (LES) describes the covariation of traits relevant for carbon and nutrient economy in different plant species. However, much less is known about the correlation of LES with leaf water economy, not only because some woody species do not follow the rules, but also because they are rarely tested on the widespread, non-native, fast-growing trees. We hypothesized that fast-growing exotic species that spread on the fast side of the LES coordinate their water-use strategies (WUS) to maintain rapid growth, and that the pattern of coordination differs between evergreen and deciduous forests. Using 4 exotic and 4 native species from evergreen and deciduous broadleaf forests in China, we measured 17 traits of LES and WUS and analyzed their functional roles in different species groups. Our results suggest that LES plays a more important role in the coexistence of species within a community, while WUS contributes more to the distribution of species across different regions. The multidimensional coordination of LES and WUS could better explain the growth and distribution of different plant species and shed light on the coexistence of species from different forest types, especially fast-growing woody exotics.

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.

Leaf habit and plant architecture integrate whole-plant economics and contextualize trait-climate associations within ecologically diverse genus Rhododendron.

Plant resource strategies negotiate a trade-off between fast growth and stress resistance, characterized by specific leaf area (SLA). How SLA relates to leaf structure and function or plant climate associations remains open for debate, and leaf habit and plant architecture may alter the costs versus benefits of individual traits. We used phylogenetic canonical correspondence analysis and phylogenetic least squares to understand the relationship of anatomy and gas exchange to published data on root, wood, architectural and leaf economics traits and climate. Leaf anatomy was structured by leaf habit and carbon to nitrogen ratio was a better predictor of gas exchange than SLA. We found significant correspondence of leaf anatomy with branch architecture and wood traits, gas exchange corresponded with climate, while leaf economics corresponded with climate, architecture, wood and root traits. Species from the most seasonal climates had the highest trait-climate correspondence, and different aspects of economics and anatomy reflected leaf carbon uptake versus water use. Our study using phylogenetic comparative methods including plant architecture and leaf habit provides insight into the mechanism of whole-plant functional coordination and contextualizes individual traits in relation to climate, demonstrating the evolutionary and ecological relevance of trait-trait correlations within a genus with high biodiversity.

Grassland sensitivity to drought is related to functional composition across East Asia and North America.

Plant traits can be helpful for understanding grassland ecosystem responses to climate extremes, such as severe drought. However, intercontinental comparisons of how drought affects plant functional traits and ecosystem functioning are rare. The Extreme Drought in Grasslands experiment (EDGE) was established across the major grassland types in East Asia and North America (six sites on each continent) to measure variability in grassland ecosystem sensitivity to extreme, prolonged drought. At all sites, we quantified community-weighted mean functional composition and functional diversity of two leaf economic traits, specific leaf area and leaf nitrogen content, in response to drought. We found that experimental drought significantly increased community-weighted means of specific leaf area and leaf nitrogen content at all North American sites and at the wetter East Asian sites, but drought decreased community-weighted means of these traits at moderate to dry East Asian sites. Drought significantly decreased functional richness but increased functional evenness and dispersion at most East Asian and North American sites. Ecosystem drought sensitivity (percentage reduction in aboveground net primary productivity) positively correlated with community-weighted means of specific leaf area and leaf nitrogen content and negatively correlated with functional diversity (i.e., richness) on an intercontinental scale, but results differed within regions. These findings highlight both broad generalities but also unique responses to drought of community-weighted trait means as well as their functional diversity across grassland ecosystems.

Leaf phosphorus fractions vary with leaf economic traits among 35 Australian woody species.

Adaptations of plants to phosphorus (P) deficiency include reduced investment of leaf P in storage (orthophosphates in vacuoles), nucleic acids and membrane lipids. Yet, it is unclear how these adaptations are associated with plant ecological strategies. Five leaf P fractions (orthophosphate P, Pi ; metabolite P, PM ; nucleic acid P, PN ; lipid P, PL ; and residual P, PR ) were analysed alongside leaf economic traits among 35 Australian woody species from three habitats: one a high-P basalt-derived soil and two low-P sandstone-derived soils, one undisturbed and one disturbed by human activities with artificial P inputs. Species at the undisturbed low-P site generally exhibited lower concentrations of total leaf P ([Ptotal ]), primarily associated with lower concentrations of Pi , and PN . The relative allocation of P to each fraction varied little among sites, except that higher PL per [Ptotal ] (rPL ) was recorded at the undisturbed low-P site than at the high-P site. This higher rPL , reflecting relative allocation to membranes, was primarily associated with lower concentrations of leaf nitrogen at the undisturbed low-P site than at the high-P site. Associations between leaf P fractions and leaf nitrogen may provide a basis for understanding the variation in plant ecological strategies dependent on soil P availability.

Coupling of leaf economic and hydraulic traits of desert plants of different life forms in the Ebinur Lake Basin

The functional traits of leaves and their internal trade-offs have been the focus of ecologists as a powerful basis for revealing plant physiology and function, and ecological strategies. The consistency of the coupled relationship between leaf economic traits and hydraulic traits among trees, shrubs, and herbs in similar ecological environments has seldom been systematically explored. We explored coupling in desert plants of different life forms and their response to the soil environment in two tree species (Populus euphratica Oliv. and Haloxylon ammodendron (C. A. Mey.) Bunge)), the shrub species (Nitraria tangutorum Bobrov and Halimodendron halodendron (Pall.) Dum. Cours.), and herbaceous species (Alhagi sparsifolia Shap. and Apocynum venetum L.) in the transition zone of the desert in the Ebinur Lake Basin, Xinjiang, China. Leaf and soil samples were collected to determine plant leaf economic traits, hydraulic traits, and soil factors. Regarding leaf economic traits, trees had higher in leaf mass per area and leaf thickness (LT). The leaf nitrogen and phosphorus contents were higher in shurbs than those in trees and herbs. Regarding hydraulic traits, shrubs had higher palisade parenchyma thickness (PT) and vein density (VD), whereas herbs had higher stomatal length (SL), spongy parenchyma thickness (ST), and leaf water potential (Ψ). In herbs, SL, ST, and Ψ had advantages, whereas PT/ST was the lowest. This study revealed an association between leaf economic and hydraulic traits in plants of different life forms in a similar ecological environment and provides a scientific basis for plant conservation strategies and ecosystem services value assessment and use in arid zones.

Quantifying foliar trait variation and covariation in sun and shade leaves using leaf spectroscopy in eastern North America

Characterizing foliar trait variation in sun and shade leaves can provide insights into inter- and intra-species resource use strategies and plant response to environmental change. However, datasets with records of multiple foliar traits from the same individual and including shade leaves are sparse, which limits our ability to investigate trait-trait, trait-environment relationships and trait coordination in both sun and shade leaves. We presented a comprehensive dataset of 15 foliar traits from sun and shade leaves sampled with leaf spectroscopy, including 424 individuals of 110 plant species from 19 sites across eastern North America. We investigated trait variation, covariation, scaling relationships with leaf mass, and the effects of environment, canopy position, and taxonomy on trait expression. Generally, sun leaves had higher leaf mass per area, nonstructural carbohydrates and total phenolics, lower mass-based chlorophyll a + b, carotenoids, phosphorus, and potassium, but exhibited species-specific characteristics. Covariation between sun and shade leaf traits, and trait-environment relationships were overall consistent across species. The main dimensions of foliar trait variation in seed plants were revealed including leaf economics traits, photosynthetic pigments, defense, and structural traits. Taxonomy and canopy position collectively explained most of the foliar trait variation. This study highlights the importance of including intra-individual and intra-specific trait variation to improve our understanding of ecosystem functions. Our findings have implications for efficient field sampling, and trait mapping with remote sensing.

Not all sweetness and light: Non‐structural carbohydrate storage capacity in tree stems is decoupled from leaf but not from root economics

ABSTRACT Non‐structural carbohydrates (NSC) are considered as indicators of the balance between tree carbon sources and sinks and reflect functional strategies throughout different biomes. However, little is known about the contribution of NSC to tree economics, and in particular, whether leaf, stem and coarse root traits co‐ordinate together into a whole‐tree economics spectrum. Twenty‐four functional traits (including NSC content) were measured in leaves, stem and coarse root xylem of up to 90 angiosperm tree species in temperate, Mediterranean and tropical climates. By performing principal component analysis and standardized major axis regression, we explored the relationships between NSC and other functional traits, as well as the effects of climate and phylogeny on these relationships. Our results revealed a covariation between leaf and coarse root NSC content and leaf economic traits, whereas stem NSC content was largely decoupled from the leaf economics spectrum that was mostly driven by leaf nitrogen content and leaf mass per area. Coarse root xylem traits were closely correlated with leaf traits, while most stem xylem traits were independent from the leaf economics spectrum but covaried with coarse root traits. Trade‐offs among traits in tropical species opposed those from other climates. Evolutionary history affected relationships among certain traits but did not change overall patterns. We conclude that due to leaf habit, an extended growing season and heightened defences, tropical species form distinct conservative resource acquisition strategies. Across all climates, as the stem must provide an efficient transport route between roots and leaves, while maintaining the display of branch and leaf organs, stem xylem design and NSC storage capacity lead to a stem economics spectrum that is largely independent from the leaf spectrum, but is coupled with that of coarse roots due to anatomical continuity. Read the free Plain Language Summary for this article on the Journal blog.

Wood density and leaf size jointly predict woody plant growth rates across (but not within) species along a steep precipitation gradient

Abstract Functional traits have been proposed to define key dimensions of plant ecological strategies, but we lack consensus on whether traits can accurately predict plant demography. Despite theoretical expectations, it has been challenging to find consistent relationships between functional traits and growth. In this study, we quantified inter‐ and intraspecific trait variation (ITV) and individual growth rates of woody plants across a steep moisture gradient that varies 10‐fold in annual precipitation (350–3700 mm) in southern Chile and used a hierarchical Bayesian model to predict growth as a function of trait values. We show that large‐leaved species with lower stem tissue density exhibited the fastest growth rates, and these two traits exhibited the highest proportion of interspecific variation. Predictions of growth improved considerably (R2 of the best model increased from 0.28 to 0.49) when species‐level multiple traits and their interactions were considered. The inclusion of ITV, however, did not improve models of growth rate. We found that trait–growth rate relationships were not always consistent across levels of biological organization; relationships observed at the interspecific level did not necessarily hold at the intraspecific level. We found that the relationships between wood density or leaf size and growth were consistent in direction across the precipitation gradient, and the relationships between leaf economics traits and growth were weak and site‐specific. Synthesis. Although using more than one functional trait considerably improved growth predictions, wood density and leaf size successfully predicted growth rates across (not within) species, which is consistent with a whole‐plant carbon economy. We assert that these two traits are intimately linked and ultimately describe a continuum of plant architecture and carbon economy that covers multiple trait syndromes.

Responses of economic and anatomical leaf traits to soil fertility factors in eight coexisting broadleaf species in temperate forests

The multidimensionality of leaf traits allows plants to have diverse survival strategies to adapt to complex living environments. Whether the anatomical traits of leaves are associated with leaf economic traits and which group of traits are more strongly correlated with soil fertility factors remains unclear. We measured four leaf economic traits, four anatomical traits, and five soil fertility factors of eight coexisting broadleaf species distributed in mixed broadleaved-Korean pine (Pinus koraiensis) forests located in Northeast China. Results show a strong interdependence between economic and anatomical traits (p < 0.05). The range of variation between economic and anatomical traits were almost equal, but the causes of variation were different. Specific leaf area was positively correlated with the abaxial epidermis, negatively correlated with the ratio of spongy tissue to leaf thickness (ST/LT), and not correlated with adaxial epidermis. Leaf dry matter content was negatively correlated with the abaxial epidermis and adaxial epidermis, positively correlated with ST/LT. Specific leaf area, palisade tissue, and ST/LT showed stronger correlation with soil fertility factors than other traits. Soil fertility factors dominating trait variation were dependent upon the trait. Our results suggest anatomical traits can be considered in economic trait dimension. The coupled relationship between anatomical and economic traits is potentially a cost-effective adaptation strategy for species to improve efficiency in resource utilization. Our results provide evidence for the complex soil-trait relationship and suggest that future studies should emphasize the role of anatomic traits in predicting soil fertility changes.

Integrating defense and leaf economic spectrum traits in a tropical savanna plant.

Allocation to plant defense traits likely depends on resource supply, herbivory, and other plant functional traits such as the leaf economic spectrum (LES) traits. Yet, attempts to integrate defense and resource acquisitive traits remain elusive. We assessed intraspecific covariation between different defense and LES traits in a widely distributed tropical savanna herb, Solanum incanum, a unique model species for studying allocations to physical, chemical, and structural defenses to mammalian herbivory. We found that in a multivariate trait space, the structural defenses - lignin and cellulose - were positively related to the resource conservative traits - low SLA and low leaf N. Phenolic content, a chemical defense, was positively associated with resource acquisitive traits - high SLA and high leaf N - while also being associated with an independent third component axis. Both principal components 1 and 3 were not associated with resource supply and herbivory intensity. In contrast, spine density - a physical defense - was orthogonal to the LES axis and positively associated with soil P and herbivory intensity. These results suggest a hypothesized "pyramid" of trade-offs in allocation to defense along the LES and herbivory intensity axes. Therefore, future attempts to integrate defense traits with the broader plant functional trait framework, such as the LES, needs a multifaceted approach that accounts for unique influences of resource acquisitive traits and herbivory risk.

Coordination in functional traits of Populus euphratica leaves under salt stress

Leaf functional traits allow plant survival and maintain their ecosystem function. Salinity affects leaf functional traits, but coordination among leaf functional traits is poorly known and may depend on salt severity. To increase our understanding of the coordination of leaf functional traits under salt stress, we determined hydraulic, gas exchange, and physiological and biochemical parameters in Populus euphratica Oliv. (P. euphratica) grown under salinity treatments, as well as gas exchange parameters under different CO2 concentrations. We found that P. euphratica can reinforce its hydraulic capacity by increasing the water transfer efficiency of both its leaves and stems when a salinity threat occurs for a specific duration of stress. Its stems were more adaptable than leaves. The economic and hydraulic traits of P. euphratica leaves were consistent during the middle stages of salt stress, but inconsistent during the onset and late stages of salt stress. There was almost no biochemical limitation under severe salinity conditions, and CO2 enrichment of P. euphratica had a greater effect on leaf economic traits. The mechanism of toxic ion exclusion based on water availability and intracellular mechanisms in leaves contributed to salt tolerance when P. euphratica was exposed to salinity stress. There was also a coordination mechanism for the plants during increasing salt stress. The leaf intracellular traits of P. euphratica can coordinate with the leaf economic and hydraulic traits and form a defense mechanism to reduce salt damage and guarantee growth under saline conditions. In conclusion, P. euphratica, the main constructional species of riparian forests, adapts to saline environments by adjustment and coordination of leaf functional traits, ensuring survival. These results provide a scientific basis for riparian forest restoration.

A survey of leaf phosphorus fractions and leaf economic traits among 12 co-occurring woody species on phosphorus-impoverished soils

Background and AimsThe leaf economic spectrum (LES) is related to dry mass and nutrient investments towards photosynthetic processes and leaf structures, and to the duration of returns on those investments (leaf lifespan, LL). Phosphorus (P) is a key limiting nutrient for plant growth, yet it is unclear how the allocation of leaf P among different functions is coordinated with the LES. We addressed this question among 12 evergreen woody species co-occurring on P-impoverished soils in south-eastern Australia.MethodsLeaf ‘economic’ traits, including LL, leaf mass per area (LMA), light-saturated net photosynthetic rate per mass (Amass), dark respiration rate, P concentration ([Ptotal]), nitrogen concentration, and P resorption, were measured for three pioneer and nine non-pioneer species. Leaf P was separated into five functional fractions: orthophosphate P (Pi), metabolite P (PM), nucleic acid P (PN), lipid P (PL), and residual P (PR; phosphorylated proteins and unidentified compounds that contain P).ResultsLL was negatively correlated with Amass and positively correlated with LMA, representing the LES. Pioneers occurred towards the short-LL end of the spectrum and exhibited higher [Ptotal] than non-pioneer species, primarily associated with higher concentrations of Pi, PN and PL. There were no significant correlations between leaf P fractions and LL or LMA, while Amass was positively correlated with the concentration of PR.ConclusionsAllocation of leaf P to different fractions varied substantially among species. This variation was partially associated with the LES, which may provide a mechanism underlying co-occurrence of species with different ecological strategies under P limitation.

Soil water content and nitrogen differentially correlate with multidimensional leaf traits of two temperate broadleaf species

The variation and correlation of leaf economics and vein traits are crucial for predicting plant ecological strategies under different environmental changes. However, correlations between these two suites of traits and abiotic factors such as soil water and nitrogen content remain ambiguous. We measured leaf economics and vein traits as well as soil water and nitrogen content for two different shade-tolerant species (Betula platyphylla and Acer mono) in four mixed broadleaved-Korean pine (Pinus koraiensis) forests along a latitudinal gradient in Northeast China. We found that leaf economics traits and vein traits were decoupled in shade-intolerant species, Betula platphylla, but significantly coupled in a shade-tolerant species, A. mono. We found stronger correlations among leaf traits in the shade tolerant species than in the shade intolerant species. Furthermore, leaf economic traits were positively correlated with the soil water gradient for both species, whereas vein traits were positively correlated with soil water gradient for the shade intolerant species but negatively correlated in the shade tolerant species. Although economic traits were positively correlated with soil nitrogen gradient in shade intolerant species but not correlated in shade tolerant species, vein traits were negatively correlated with soil nitrogen gradient in shade tolerant species but not correlated in shade intolerant species. Our study provides evidence for distinct correlations between leaf economics and vein traits and local abiotic factors of species differing in light demands. We recommend that the ecological significance of shade tolerance be considered for species when evaluating ecosystem functions and predicting plant responses to environmental changes.

Vein Network and Climatic Factors Predict the Leaf Economic Spectrum of Desert Plants in Xinjiang, China.

The leaf economic spectrum (LES) has been repeatedly verified with regional and global datasets. However, the LES of desert plants and its drivers has not been fully explored at the species level. In this study, we sampled three desert perennial plant species (Alhagi sparsifolia, Karelinia caspia, and Apocynum venetum) at three different geographical areas of distribution in Xinjiang, China, and measured 10 leaf economic traits to determine their strategy of resource utilization. The scores of the first axis from the principal component analysis of 10 leaf economic traits as a continuous variable define the LES. This study showed that the LES did exist in desert plants in this region. The leaf economic spectrum shifted from a more conservative strategy to a more acquisitive strategy with increasing contents of soil potassium (K) and the ratio of K to phosphorus. Except for the vein density of A. venetum, which quadratically correlated with LES, the vein density, distance between veins, and vein loopiness significantly positively correlated with the LES (p < 0.05), indicating a covariation and tradeoff relationship. The annual mean temperature was significantly negatively correlated with LES, while the annual mean precipitation (MAP) and the aridity index (AI), which was calculated by the ratio of MAP to potential evapotranspiration, significantly positively correlated with the LES. Of these, vein loopiness and AI were more effective at predicting the change in LES from anatomical and climatic perspectives owing to their high regression coefficients (R2). The findings of this study will substantially improve the understanding of the strategies of desert plants to utilize resources and predict the structure and function of ecosystems.

Demographic trade-offs and functional shifts in a hurricane-impacted tropical forest.

Understanding shifts in the demographic and functional composition of forests after major natural disturbances has become increasingly relevant given the accelerating rates of climate change and elevated frequency of natural disturbances. Although plant demographic strategies are often described across a slow-fast continuum, severe and frequent disturbance events influencing demographic processes may alter the demographic trade-offs and the functional composition of forests. We examined demographic trade-offs and the shifts in functional traits in a hurricane-disturbed forest using long-term data from the Luquillo Forest Dynamics Plot (LFPD) in Puerto Rico. We analysed information on growth, survival, seed rain and seedling recruitment for 30 woody species in the LFDP. In addition, we compiled data on leaf, seed and wood functional traits that capture the main ecological strategies for plants. We used this information to identify the main axes of demographic variation for this forest community and evaluate shifts in community-weighted means for traits from 2000 to 2016. The previously identified growth-survival trade-off was not observed. Instead, we identified a fecundity-growth trade-off and an axis representing seedling-to-adult survival. Both axes formed dimensions independent of resprouting ability. Also, changes in tree species composition during the post-hurricane period reflected a directional shift from seedling and tree communities dominated by acquisitive towards conservative leaf economics traits and large seed mass. Wood specific gravity, however, did not show significant directional changes over time. Our study demonstrates that tree demographic strategies coping with frequent storms and hurricane disturbances deviate from strategies typically observed in undisturbed forests, yet the shifts in functional composition still conform to the expected changes from acquisitive to conservative resource-uptake strategies expected over succession. In the face of increased rates of natural and anthropogenic disturbance in tropical regions, our results anticipate shifts in species demographic trade-offs and different functional dimensions.

Inter- and Intraspecific Variation Patterns of Leaf Phenotypic Traits in Two Sympatric Pine Species

Individual persistence under changing climate conditions can be aided by phenotypic plasticity. Needle morphology reflects pine species’ adaptation to their habitats, and adaptive plasticity is beneficial to pine survival. As two closely related pines, Pinusmassoniana Lamb. and Pinus hwangshanensis W. Y. Hsia sympatrically occur in the subtropical region of China, forming hilly forests (i.e., subtropical evergreen needleleaf forests) at lower elevations and montane forests (i.e., temperate evergreen needleleaf forests) at medium and high elevations. However, little is known about the intraspecific phenotypic variation patterns of the two pine species and their relationship with environmental factors. Here, we investigated the interspecific and intraspecific variation patterns of needle traits in the two sympatric pine species, focusing on six traits—i.e., leaf length, leaf thickness, leaf width, leaf area, specific leaf area, and leaf dry matter content—as well as the relationship between needle traits and environmental factors. We found significant phenotypic trait differences among populations of each species. Except for leaf length and leaf width, variation within species was greater than variation between species in the needle traits measured. Even more intriguingly, the leaf size traits (i.e., length, width, thickness) were more conservative than the leaf economic traits (i.e., specific leaf area and leaf dry matter content). In other words, the intraspecific variability of the former was weaker than that of the latter. The nature of P. massoniana needle traits was mainly shaped by latitude, while the needle traits of P. hwangshanensis were significantly affected by annual precipitation. Therefore, phenotypic plasticity may be an essential mechanism for the two pine species to better cope with changing external conditions. The intraspecific variation patterns found in the two pine species and the relationships between traits and environmental factors can provide substantial scientific data for large-scale exploration of intraspecific phenotypic variation in pine species and their breeding practice.

Linking leaf economic and hydraulic traits with early-age growth performance and survival of Eucalyptus pauciflora.

Selection on plant functional traits may occur through their direct effects on fitness (or a fitness component), or may be mediated by attributes of plant performance which have a direct impact on fitness. Understanding this link is particularly challenging for long-lived organisms, such as forest trees, where lifetime fitness assessments are rarely achievable, and performance features and fitness components are usually quantified from early-life history stages. Accordingly, we studied a cohort of trees from multiple populations of Eucalyptus pauciflora grown in a common-garden field trial established at the hot and dry end of the species distribution on the island of Tasmania, Australia. We related the within-population variation in leaf economic (leaf thickness, leaf area and leaf density) and hydraulic (stomatal density, stomatal length and vein density) traits, measured from two-year-old plants, to two-year growth performance (height and stem diameter) and to a fitness component (seven-year survival). When performance-trait relationships were modelled for all traits simultaneously, statistical support for direct effects on growth performance was only observed for leaf thickness and leaf density. Performance-based estimators of directional selection indicated that individuals with reduced leaf thickness and increased leaf density were favoured. Survival-performance relationships were consistent with size-dependent mortality, with fitness-based selection gradients estimated for performance measures providing evidence for directional selection favouring individuals with faster growth. There was no statistical support for an effect associated with the fitness-based quadratic selection gradient estimated for growth performance. Conditional on a performance measure, fitness-based directional selection gradients estimated for the leaf traits did not provide statistical support for direct effects of the focal traits on tree survival. This suggested that, under the environmental conditions of the trial site and time period covered in the current study, early-stage selection on the studied leaf traits may be mediated by their effects on growth performance, which in turn has a positive direct influence on later-age survival. We discuss the potential mechanistic basis of the direct effects of the focal leaf traits on tree growth, and the relevance of a putative causal pathway of trait effects on fitness through mediation by growth performance in the studied hot and dry environment.