Leaf Economics Research Articles

Leaf mass per area: An investigation into the application of the ubiquitous functional trait from a paleobotanical perspective.

Leaf mass per area (LMA) is a widely used functional trait in both neobotanical and paleobotanical research that provides a window into how plants interact with their environment. Paleobotanists have used site-level measures of LMA as a proxy for climate, biome, deciduousness, and community-scale plant strategy, yet many of these relationships have not been grounded in modern data. In this study, we evaluated LMA from the paleobotanical perspective, seeking to add modern context to paleobotanical interpretations and discover what a combined modern and fossil data set can tell us about how LMA can be best applied toward interpreting plant communities. We built a modern data set by pulling plant trait data from the TRY database, and a fossil data set by compiling data from studies that have used the petiole-width proxy for LMA. We then investigated the relationships of species-mean, site-mean, and site-distribution LMA with different climatic, phylogenetic, and physiognomic variables. We found that LMA distributions are correlated with climate, site taxonomic composition, and deciduousness. However, the relative contributions of these factors are not distinctive, and ultimately, LMA distributions cannot accurately reconstruct the biome or climate of an individual site. The correlations that make up the leaf economics spectrum are stronger than the correlations between LMA and climate, phylogeny, morphospace, or depositional environment. Fossil LMA should be understood as the culmination of the influences of these variables rather than as a predictor.

Foliar methane and nitrous oxide fluxes in tropical tree species

Methane (CH₄) and nitrous oxide (N₂O) are critical biogenic greenhouse gases (GHGs) with global warming potentials substantially greater than that of carbon dioxide (CO₂). The exchange of these gases in tropical forests, particularly via foliar processes, remains poorly understood. We quantified foliar CH₄ and N₂O fluxes among tropical tree species and examined their potential association with the leaf economics spectrum (LES) traits. Sampling within Lawachara National Park, Bangladesh, we used in-situ measurements of foliar CH₄ and N₂O fluxes employing off-axis integrated cavity output spectroscopy (CH₄, CO₂ and H₂O) and optical feedback–cavity enhanced absorption spectroscopy (N₂O) analyzers. Leaves were measured under dark, low, and high (0, 100, and 1000 μmol·m−2·s−1) light conditions. Surveyed tree species exhibited both net foliar uptake and efflux of CH₄, with a mean flux not different from zero, suggesting negligible net foliar emissions at the stand level. Plant families showed differences in CH₄, but not N₂O fluxes. Consistent efflux was observed for N₂O, with a mean of 0.562 ± 0.060 pmol·m−2·s−1. Pioneer species exhibited a higher mean N₂O flux (0.81 ± 0.17 pmol·m−2·s−1) compared to late-successional species (0.37 ± 0.05 pmol·m−2·s−1). Pioneer species also showed a trend toward a higher mean CH₄ flux (0.24 ± 0.21 nmol·m−2·s−1) compared to mid-successional (−0.01 ± 0.26 nmol·m−2·s−1) and late-successional species (−0.05 ± 0.28 nmol·m−2·s−1). Moreover, among all leaf traits within the leaf economic spectrum, a significant positive relationship was observed between leaf N₂O flux and total leaf nitrogen. Our results suggest that pioneer tree species significantly contribute to net CH₄ and N₂O emissions, potentially counteracting the carbon sequestration benefits in regenerating tropical forests. These findings indicate that accurate GHG budgeting should include direct measurements of foliar CH₄ and N₂O fluxes. Moreover, the results suggest that forest conservation and management strategies that prioritize late successional species will better mitigate GHG emissions.

Measuring standardized functional leaf traits of aquatic carnivorous plants – challenges and opportunities

Aquatic carnivorous plants (ACP) are an important component of humic nutrient-poor freshwater environments. However, these habitats are facing multiple impacts that ultimately lead to habitat degradation and declining ACP populations. Functional traits, particularly those within the leaf economics spectrum, are a valuable tool for studying plant adaptation strategies and plasticity. Given their unique morphological structure, ACP are essentially excluded from functional comparisons, which potentially limits our knowledge about the ecological roles of these species compared to non-carnivorous ones. In this study, we developed a protocol for measuring the leaf functional traits of ACP (leaf fresh and dry weight, leaf area, leaf dry matter content, specific leaf area, leaf pigment content, leaf phosphorus, nitrogen and carbon contents), and carnivory-related traits (number of traps and investment in carnivory). We measured 15 traits in seven ACP species (Aldrovanda vesiculosa, Utricularia australis, U. bremii, U. intermedia, U. ochroleuca, U. stygia, U. vulgaris), grown in the outdoor collection of aquatic and wetland plants of the Institute of Botany CAS at Třeboň, the Czech Republic. We used the functional traits of other macrophyte groups/species (lemnids, Nuphar lutea, Ceratophyllum demersum), collected with a similar methodology, to assess the comparability of ACP traits. We identified the “functional unit”, a modular structure, including one leaf node, plus an internode, which performs the function of a leaf in ACP, and selected its position along the stem to reflect species-specific growth rates. We collected 714 new trait records for the target ACP. Based on a multivariate trait space representation (PCA), ACP were distinct from the other macrophyte groups/species, which highlights these species’ structural and physiological peculiarities. Nonetheless, ACP entirely overlapped the comparison data along the first PCA axis, and most of the traits lay within the ranges observed for other macrophyte groups/species, which demonstrates the comparability of the ACP traits measured by the new protocol. Applying this protocol in ecological studies could shed light on the adaptations of ACP to environmental variability, with important conservation implications.

Global patterns of plant functional traits and their relationships to climate

Plant functional traits (FTs) determine growth, reproduction and survival strategies of plants adapted to their growth environment. Exploring global geographic patterns of FTs, their covariation and their relationships to climate are necessary steps towards better-founded predictions of how global environmental change will affect ecosystem composition. We compile an extensive global dataset for 16 FTs and characterise trait-trait and trait-climate relationships separately within non-woody, woody deciduous and woody evergreen plant groups, using multivariate analysis and generalised additive models (GAMs). Among the six major FTs considered, two dominant trait dimensions—representing plant size and the leaf economics spectrum (LES) respectively—are identified within all three groups. Size traits (plant height, diaspore mass) however are generally higher in warmer climates, while LES traits (leaf mass and nitrogen per area) are higher in drier climates. Larger leaves are associated principally with warmer winters in woody evergreens, but with wetter climates in non-woody plants. GAM-simulated global patterns for all 16 FTs explain up to three-quarters of global trait variation. Global maps obtained by upscaling GAMs are broadly in agreement with iNaturalist citizen-science FT data. This analysis contributes to the foundations for global trait-based ecosystem modelling by demonstrating universal relationships between FTs and climate.

Fine Root Traits across Different Root Orders and Their Associations with Leaf Traits in 15 Co-Occurring Plant Species from the Desert-Oasis Transition Zone in the Hexi Corridor, Gansu Province, China.

Fine root traits embody trade-offs between resource acquisition and conservation in plants. Yet, the differentiation of these traits across root orders, the existence of a root economics spectrum (RES) spanning these orders, and their linkage with leaf traits remain underexplored. In this study, we analyzed the first three root orders and leaf traits of 15 co-occurring plant species, including ten herbs and five shrubs, from the desert-oasis transition zone of the Hexi Corridor. We measured twelve morphological and chemical traits to investigate the relationships between root and leaf traits. Our results revealed significant variation in root traits both among species and within species across different root orders. We identified RES that spanned root orders, with higher-order roots exhibiting more conservative traits and lower-order roots displaying traits aligned with resource acquisition. Additionally, leaf and fine root traits showed partially decoupled adaptive strategies, yet evidence also supported the existence of a leaf economics spectrum (LES) and a potentially two-dimensional whole plant economics spectrum (WPES). Our findings suggest synergistic resource allocation strategies between fine roots and the entire plant, emphasizing the importance of root order in understanding fine root structure, function, and their interactions with other plant organs. These insights advance the understanding of fine root traits and their integration within the broader plant economics spectrum. Nevertheless, the differences in fine root traits across root orders, the presence of a root economics spectrum (RES) spanning these orders, and the relationships between fine root and leaf traits remain underexplored. We examined the first three root orders and leaves of 15 co-occurring plant species (ten herbs and five shrubs) from the desert-oasis transition zone in the Hexi Corridor, measured twelve key morphological and chemical traits. We observed substantial variation in root traits among species and root orders within species. The root economics spectrum (RES) spanned across root orders, with higher-order roots positioned at the conservative end and lower-order roots at the acquisitive end of the "investment-return" strategy axis. Leaf and fine root traits of the 15 co-occurring plant species exhibited partially decoupled adaptive strategies. However, there was also evidence for the presence of a leaf economics spectrum (LES) and a whole plant economics spectrum (WPES), with the WPES potentially being two-dimensional. Furthermore, our findings suggest synergistic resource strategies between fine roots and the whole plant. Concurrently, the significant interspecific and intraspecific differences in fine root traits, combined with the presence of a root economics spectrum across root orders, underscore the critical importance of root order in studying fine root structure, function, and their associations with other plant organs. Our findings offer valuable insights for future research on fine root traits, the RES, and their integration with the whole plant economics spectrum.

Unfolding the leaf economics spectrum for wheat: Trait analysis and genomic associations across cultivars

The leaf economics spectrum (LES) is an ecophysiological concept that describes the trade-offs between leaf structural and physiological traits. It has been extensively studied across various scales. However, the coordination hypothesis has rarely been tested at the intraspecific scale, especially in crops, for understanding yield increases or predicting evolutionary trajectories. Here, we first tested the relationships among leaf traits and examined the genetic coordination among 209 wheat genotypes. Compared to non-crop grass species, wheat is a fast-growing species, and tends to have a higher value of photosynthetic rate, leaf nitrogen concentration and leaf respiration rate at a given leaf mass per area, although it does align with the predicted direction of the “fast-slow” spectrum. We conducted a principal component analysis (PCA) to compare different traits within wheat. The first axis from PCA (ranging from slow to fast of plant economic investment) is significantly positively associated with the agronomic traits, especially grain yield (R2=0.11, P<0.001). Partially independent changes in leaf nitrogen content and leaf mass per area may allow crops to maximize photosynthetic rates without sacrificing leaf lifespan. The results reveal that some loci are simultaneously associated with different traits, which may be the genetic basis for the formation of trait-trait relationships. The current study deepens the understanding of LES traits in wheat at the intraspecific and genetic levels, supporting the trait-based adaptation strategies to improve wheat productivity and resource-use efficiency.

Life at the conservative end of the leaf economics spectrum: intergeneric variation in the allocation of phosphorus to biochemical fractions in species of Banksia (Proteaceae) and Hakea (Proteaceae).

In severely phosphorus (P)-impoverished environments, plants have evolved to use P very efficiently. Yet, it is unclear how P allocation in leaves contributes to their photosynthetic P-use efficiency (PPUE) and position along the leaf economics spectrum (LES). We address this question in 10 species of Banksia and Hakea, two highly P-efficient Proteaceae genera. We characterised traits in leaves of Banksia and Hakea associated with the LES: leaf mass per area, light-saturated photosynthetic rates, P and nitrogen concentrations, and PPUE. We also determined leaf P partitioning to five biochemical fractions (lipid, nucleic acid, metabolite, inorganic and residual P) and their possible association with the LES. For both genera, PPUE was negatively correlated with fractional allocation of P to lipids, but positively correlated with that to metabolites. For Banksia only, PPUE was negatively correlated with residual P, highlighting a strategy contrasting to that of Hakea. Phosphorus-allocation patterns significantly explained PPUE but were not linked to the resource acquisition vs resource conservation gradient defined by the LES. We conclude that distinct P-allocation patterns enable species from different genera to achieve high PPUE and discuss the implications of different P investments. We surmise that different LES axes representing different ecological strategies coexist in extremely P-impoverished environments.

Variations in the leaf economics spectrum, anatomical, ultrastructural, and stomatal traits of five tree species in the urban-rural air pollution environment

Rapid urbanization has contributed to global increases in air pollution derived from urban areas. Unlike natural forests, urban forests are exposed to higher concentrations of airborne pollutants due to the strong urban-suburban-rural pollutant emission gradients. However, there remains a pressing lack of available information pertaining to the urban air pollution-related effects on the leaf economics spectrum, anatomical, ultrastructural, and stomatal traits of tree species along an urban-rural gradient. Here, the degree to which urban air pollution impacts the adaption of greening tree species and associated service functions was assessed by sampling five common tree species (Acer pictum, Fraxinus chinensis, Koelreuteria paniculata, Salix babylonica, Sophora japonica) along urban-rural-natural forests in the Beijing metropolitan region of China. These analyses revealed a significant reduction in leaf mass per unit area (–13.4 %), leaf thickness (–16.7 %), and stomatal area (–27.5 %) with increasing proximity to areas of greater air pollution that coincide with significant increases in leaf tissue density (+12.6 %), leaf nitrogen content (+10.1 %), relative chlorophyll content (+2.7 %), and stomatal density (+11.9 %). Higher air pollution levels were associated with organelle changes including gradual disintegration of chloroplasts, larger intercellular spaces and apparent starch and plastoglobuli deposition. Air pollution was conducive to the strengthening of the trade-off potential and adaptation strategies of trees in urban ecosystems, which are associated with trees with a rapid investment return strategy associated with thick leaves and strong photosynthetic capacity. These results provide strong empirical evidence of the profound air pollution-induced changes in leaf functional traits and adaption ability of urban forest tree species.

Leaf economics spectrum of wheat in response to multigenerational atmospheric elevated CO2

While numerous studies have explored the impacts of elevated atmospheric CO2 concentration (e[CO2]) on leaf traits in plants within a single life cycle, the effects of multigenerational e[CO2] on leaf functional traits and their interrelationships remain obscure in major crops. Leaf economics spectrum (LES) comprises a group of leaf traits, which describes trade-offs between the leaf characteristics that reflect carbon economy or returns on the investments of nutrients and dry mass. In this study, the wheat seeds harvested from a 7-generation experiment under the e[CO2] (800 μmol L−1) or the ambient CO2 concentration were planted to assess changes in the LES. Our results revealed a consistent increase in mass-normalized leaf nitrogen and phosphorus concentration under the e[CO2] over the generations. Photosynthetic phosphorus use efficiency was positively correlated with photosynthetic rate per unit mass, dark respiration per unit mass, photosynthetic rate per unit area, leaf phosphorus per unit area, and dark respiration per unit area. Moreover, we used structural equation model to illustrate the network of relationships among LES traits and shoot dry weight, providing insights into the effects of the multigenerational e[CO2] on wheat growth. This study contributes novel perspectives to our understandings of wheat responses to long-term climate change.

Characteristics, Relationships, and Anatomical Basis of Leaf Hydraulic Traits and Economic Traits in Temperate Desert Shrub Species.

Shrubs are a key component of desert ecosystems, playing a crucial role in controlling desertification and promoting revegetation, yet their growth is often impeded by drought. Leaf hydraulic traits and economic traits are both involved in the process of water exchange for carbon dioxide. Exploring the characteristics, relationships, and anatomical basis of these two suites of traits is crucial to understanding the mechanism of desert shrubs adapting to the desert arid environment. However, the relationship between these two sets of traits currently remains ambiguous. This study explored the leaf hydraulic, economic, and anatomical traits of 19 desert shrub species. The key findings include the following: Relatively larger LT values and smaller SLA values were observed in desert shrubs, aligning with the "slow strategy" in the leaf economics spectrum. The relatively high P50leaf, low HSMleaf, negative TLPleaf, and positive HSMtlp values indicated that severe embolism occurs in the leaves during the dry season, while most species were able to maintain normal leaf expansion. This implies a "tolerance" leaf hydraulic strategy in response to arid stress. No significant relationship was observed between P50leaf and Kmax, indicating the absence of a trade-off between hydraulic efficiency and embolism resistance. Certain coupling relationships were observed between leaf hydraulic traits and economic traits, both of which were closely tied to anatomical structures. Out of all of the leaf traits, LT was the central trait of the leaf traits network. The positive correlation between C content and WPleaf and HSMleaf, as well as the positive correlation between N content and HSMtlp, suggested that the cost of leaf construction was synergistic with hydraulic safety. The negative correlation between SLA, P content, GCL, and SAI suggested a functional synergistic relationship between water use efficiency and gas exchange rate. In summary, this research revealed that the coupling relationship between leaf hydraulic traits and economic traits was one of the important physiological and ecological mechanisms of desert shrubs for adapting to desert habitats.

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.

Multifaceted leaf litter traits shape soil fauna communities: Evidence from subtropical monocultural plantations

The important role of litter traits in shaping soil fauna communities has been increasingly acknowledged. It remains uncertain how fauna community features are associated with multiple litter traits, especially over the long term. With equal-aged monocultural plantations of 20 species in a subtropical climate in Hunan, China, we tested whether fauna communities were different after 40 years due to the landuse and choice of tree species, and the extent to which such disparity can be explained by litter traits. For each plantation plus a natural forest plot, we inventoried fauna in three layers (fresh litter, fragmentation and humus), measured fresh litter traits (leaf economics, size and shape and defenses), litter quantity and soil characteristics (soil properties, root and microbial biomass). Compared with natural forests, most plantations had similar fauna diversity indexes, but markedly lower density (-53.84%). Both density and diversity indexes generally increased from litter layers to humus: 0.89–1.71 for density, 32.45–38.03 for richness, 2.46–3.01 for Shannon-Wiener index, 0.82–0.92 for Gini-Simpson index and 0.74–0.84 for Pielou's evenness, respectively. Significant correlations between soil fauna indicators and litter traits were almost absent for litter layers, but were frequently observed in the humus layer. In the humus layer, soil fauna density was associated with traits related to economics (e.g., specific leaf area and dry matter content), defenses (e.g., phenol concentration) and metal element concentrations, but not with any size and shape traits, whereas soil fauna diversity could be associated with all the above trait categories. Both the leaf economics spectrum and size and shape spectrum could be significantly associated with the fauna indicators of the humus layer. Soil fauna indicators were more closely associated with litter traits than litter quantity and soil characteristics. This study reveals a link between soil fauna communities and litter traits at both individual traits and overall spectrum levels. The results emphasize the significance of segregating litter and soil layers in soil fauna studies, and the essential role of natural forests in preserving soil biodiversity.

Variations and trade-offs in leaf and culm functional traits among 77 woody bamboo species

BackgroundWoody bamboos are the only diverse large perennial grasses in mesic-wet forests and are widely distributed in the understory and canopy. The functional trait variations and trade-offs in this taxon remain unclear due to woody bamboo syndromes (represented by lignified culm of composed internodes and nodes). Here, we examined the effects of heritable legacy and occurrence site climates on functional trait variations in leaf and culm across 77 woody bamboo species in a common garden. We explored the trade-offs among leaf functional traits, the connection between leaf nitrogen (N), phosphorus (P) concentrations and functional niche traits, and the correlation of functional traits between leaves and culms.ResultsThe Bayesian mixed models reveal that the combined effects of heritable legacy (phylogenetic distances and other evolutionary processes) and occurrence site climates accounted for 55.10–90.89% of the total variation among species for each studied trait. The standardized major axis analysis identified trade-offs among leaf functional traits in woody bamboo consistent with the global leaf economics spectrum; however, compared to non-bamboo species, the woody bamboo exhibited lower leaf mass per area but higher N, P concentrations and assimilation, dark respiration rates. The canonical correlation analysis demonstrated a positive correlation (ρ = 0.57, P-value < 0.001) between leaf N, P concentrations and morphophysiology traits. The phylogenetic principal components and trait network analyses indicated that leaf and culm traits were clustered separately, with leaf assimilation and respiration rates associated with culm ground diameter.ConclusionOur study confirms the applicability of the leaf economics spectrum and the biogeochemical niche in woody bamboo taxa, improves the understanding of woody bamboo leaf and culm functional trait variations and trade-offs, and broadens the taxonomic units considered in plant functional trait studies, which contributes to our comprehensive understanding of terrestrial forest ecosystems.

Substantial variation of the two‐dimensional spectrum of senescent leaf in subalpine forest along elevational gradients

AbstractLeaf senescence have shed light on its contribution to the recovery of nutrients and signaling of the transition from the active to the dormant stage in winter. The manner by which climatic and edaphic factors and plant phylogeny affect the senescent leaf economics spectrum (more mineral elements included) of different growth habits along elevational gradients in mountain ecosystems is highly uncertain. To fill this knowledge gap, we examined 15 representative subalpine forest species' senescent leaf functional traits (i.e., specific leaf area [SLA], leaf dry matter content [DMC], pH, carbon [C], nitrogen [N], phosphorus [P], potassium [K], calcium [Ca], sodium [Na], magnesium [Mg], manganese [Mn], aluminum [Al], iron [Fe], C:N, N:P, and C:P ratios) to determine the senescent leaf economics spectrum along elevational gradients in the Hengduan Mountains, China. Our results demonstrate that all the functional traits of senescent leaf showed strong correlations with one another, and K was observed as the hub for the leaf economics spectrum. Irrespective of the difference among growth habits (arbor vs. shrub), leaf habits (deciduous vs. evergreen), and leaf types (broad vs. needle), the variations in the two‐dimensional spectrum of the senescent leaf trait exhibited uniformly positive associations with elevation, primarily attributed to climatic drivers. However, the modulation of the senescent leaf economics spectrum by soil properties and plant phylogeny was surprisingly modest. Collectively, our results revealed that the resource trade‐off of senescent leaf showed a trend from acquisition to conservation along elevational gradients, which could broaden the plant economics spectrum to include senescent leaf.

Impact of thinning on leaf economics, plant hydraulics, and growth dynamics

Integrating climate change concerns into forest management strategies remains challenging. Among management strategies, thinning has been proven to alter major components underlying the carbon and water cycles over the short term. However, the functional adaptability of managed forests to cope with long-term drought remains unclear. This study aims to quantify the influence of thinning on key plant functional traits for two pine species, and their impacts on the fundamental processes of tree growth and transpiration. We conducted an experimental silvicultural trial with varying thinning intensities in two Pinus sylvestris and Pinus nigra plantations with the following specific objectives: i) to assess the impact of thinning on major traits involved in the leaf economic spectrum (LES), and hydraulic relations, and ii) to understand which of these traits reflect fundamental aspects of plant growth and transpiration, and their relations with environment. We used a combination of dendrochronological (basal area increment, BAI), sapflow (Fd), and tree ring isotope (δ13C) data, along with key functional and physiological traits including photosynthesis (A), leaf mass area (LMA), Huber value (Hv), sapwood density (Wd), and predawn to midday water potential (Ψpd, Ψmd).Our results revealed species-specific responses, with P. nigra exhibiting a conservative growth strategy, whereas P. sylvestris displayed increased growth after thinning. Despite thinning influenced both BAI and wood δ13C with notable species-dependent variations, we evidenced a general convergence in the relationships between LES traits and Hv across species, treatments and seasons (p < 0.01). BAI responses to thinning translated into predictable changes in key hydraulic traits and adaptive changes in physiological performance. For instance, BAI was inversely related to LMA and Hv. Consistent with LES, variations in LMA influenced A. In turn, seasonal variations in LMA scaled negatively with Ψpd, and Ψmd (p < 0.01). Such a functional adaptation suggests that thinning promotes growth while favouring forest resilience in the long term. This study provides practical implications for the implementation of a trait-based approach into silvicultural frameworks and anticipating the consequences of climate change for managed forest ecosystems.

Extremely thin but very robust: Surprising cryptogam trait combinations at the end of the leaf economics spectrum

Leaf economics spectrum (LES) describes the fundamental trade-offs between leaf structural, chemical, and physiological investments. Generally, structurally robust thick leaves with high leaf dry mass per unit area (LMA) exhibit lower photosynthetic capacity per dry mass (Amass). Paradoxically, “soft and thin-leaved” mosses and spikemosses have very low Amass, but due to minute-size foliage elements, their LMA and its components, leaf thickness (LT) and density (LD), have not been systematically estimated. Here, we characterized LES and associated traits in cryptogams in unprecedented details, covering five evolutionarily different lineages. We found that mosses and spikemosses had the lowest LMA and LT values ever measured for terrestrial plants. Across a broad range of species from different lineages, Amass and LD were negatively correlated. In contrast, Amass was only related to LMA when LMA was greater than 14 g cm−2. In fact, low Amass reflected high LD and cell wall thickness in the studied cryptogams. We conclude that evolutionarily old plant lineages attained poorly differentiated, ultrathin mesophyll by increasing LD. Across plant lineages, LD, not LMA, is the trait that represents the trade-off between leaf robustness and physiology in the LES.

Tree and mycorrhizal fungal diversity drive intraspecific and intraindividual trait variation in temperate forests: Evidence from a tree diversity experiment

Abstract The study of tree species coexistence is crucial to understand the assembly of forest communities. In this context, trees adjust their traits in response to the interactions with other trees and, specifically, as a result of the competition for resources. Further, mycorrhizal fungal diversity and associations are important drivers of ecosystem functioning in forests, but their role as drivers of intraspecific trait variation has been disregarded. Here, we studied intraspecific trait variation of trees in response to tree and mycorrhizal fungal diversity. We sampled 3200 leaves from 640 trees belonging to 10 native, deciduous species in a tree diversity experiment in Central Germany. This experiment relies on the combination of gradients of tree richness and mycorrhizal associations. To handle large amounts of leaf samples, we acquired leaf‐level spectral data and used deep learning to predict values for five leaf traits from the leaf economics spectrum (LES): specific leaf area, leaf dry matter content, carbon to nitrogen ratio, carbon content and phosphorus content. For every tree, we calculated the mean value for every trait and two multi‐trait functional indices (functional richness and functional dispersion) based on values for individual leaves. Finally, we used sequencing‐based data to assess the richness of mycorrhizal fungi associated with the trees. We found that tree and mycorrhizal fungi richness had an effect on different leaf functional traits. Specifically, tree richness positively affected specific leaf area and, additionally, had a negative effect on the functional indicies, which revealed that the phenotypic diversity within the tree crown decreased with tree species richness. In addition, leaf carbon to nitrogen ratio decreased with increasing arbuscular mycorrhizal fungal richness in both arbuscular and ectomycorrhizal tree species. Finally, we did not find differences between arbuscular and ectomycorrhizal trees regarding their location within the LES. Our results suggest that trees modify their strategy in response to local tree diversity, not only by shifting trait values but also by shifting the variability intraindividually. In addition, higher mycorrhizal fungal diversity does not seem to lead to higher complementarity, but instead, tree and mycorrhizal fungi affect different aspects of leaf traits. Read the free Plain Language Summary for this article on the Journal blog.

Contrasting coordination of non-structural carbohydrates with leaf and root economic strategies of alpine coniferous forests.

Non-structural carbohydrates (NSCs), as the labile fraction and dominant carbon currency, are essential mediators of plant adaptation to environments. However, whether and how NSC coordinates with plant economic strategy frameworks, particularly the well-recognized leaf economics spectrums (LES) and root economics space (RES), remains unclear. We examined the relationships between NSC and key plant economics traits in leaves and fine roots across 90 alpine coniferous populations on the Tibetan Plateau, China. We observed contrasting coordination of NSC with economics traits in leaves and roots. Leaf total NSC and soluble sugar aligned with the leaf economic spectrum, conveying a trade-off between growth and storage in leaves. However, NSC in roots was independent of the root economic spectrum, but highly coordinated with root foraging, with more starch and less sugar in forage-efficient, thinner roots. Further, NSC-trait coordination in leaves and roots was, respectively, driven by local temperature and precipitation. These findings highlight distinct roles of NSC in shaping the above- and belowground multidimensional economics trait space, and NSC-based carbon economics provides a mechanistic understanding of how plants adapt to heterogeneous habitats and respond to environmental changes.

A framework to study and predict functional trait syndromes using phylogenetic and environmental data

Abstract Traits do not evolve in isolation but often as part of integrated trait syndromes, yet the relative contributions of environmental effects and evolutionary history on traits and their correlations are not easily resolved. In the present study, we develop a methodological framework to elucidate eco‐evolutionary patterns in functional trait syndromes. We do so by separating the amount of variance and covariance related to phylogenetic heritage and environmental variables (environmental phylogenetic conservatism), only phylogenetic heritage (non‐attributed phylogenetic conservatism) and only to environmental variables (evolutionarily labile environmental effects). Variance–covariance structures of trait syndromes are displayed as networks. We then use this framework to guide a newly derived imputation method based on machine learning models that predict trait values for unsampled taxa, considering environmental and phylogenetic information as well as trait covariation. TrEvol is presented as an R package providing a unified set of methodologies to study and predict multivariate trait patterns and improve our capacity to impute trait values. To illustrate its use, we leverage both simulated data and species‐level traits related to hydraulics and the leaf economics spectrum, in relation to an aridity index, demonstrating that most trait correlations can be attributed to environmental phylogenetic conservatism. This conceptual framework can be employed to examine issues ranging from the evolution of trait adaptation at different phylogenetic depths to intraspecific trait variation.

Soil microenvironmental variation drives below‐ground trait variation and interacts with macroclimate to structure above‐ground trait variation of arctic shrubs

Abstract Intraspecific trait variation can influence plant performance in different environments and may thereby determine the ability of individual plants to respond to climate change. However, our understanding of its patterns and environmental drivers across different spatial scales is incomplete, especially in understudied regions like the Arctic. To fill this knowledge gap, we examined above‐ground and below‐ground traits from three shrub taxa expanding across the tundra biome and evaluated their relationships with multiple microenvironmental and macroclimatic factors. The traits reflected plant size and structure (plant height, leaf area and root to shoot ratio), leaf economics (specific leaf area, nitrogen content), and root economics and collaboration with mycorrhizal fungi (specific root length, root tissue density, nitrogen content, and ectomycorrhizal colonisation intensity). We also measured leaf and root δ15N and leaf δ13C to characterise nitrogen source and acquisition pathways and plant water stress. Traits were measured in replicated plots (N = 135) varying in soil microclimate, thaw depth and organic layer thickness established across five sites spanning a macroclimate gradient in northern Alaska. This hierarchical design allowed us to disentangle the independent and combined effects of fine‐scale and broad‐scale factors on intraspecific trait variation. We found substantial intraspecific variation at fine spatial scales for most traits and less variation along the macroclimate gradient and between shrub taxa. Consistent with these patterns, microenvironmental factors, mainly soil moisture and thaw depth, interacted with macroclimate, mainly climatic water deficit, to structure size‐structural and leaf trait variation. In contrast, most root traits responded additively to thaw depth and macroclimate. Synthesis. Our results demonstrate that above‐ground and below‐ground tundra shrub traits respond differently to microenvironmental and macroclimatic variation. These differing responses contribute to substantial trait variation at fine spatial scales and may decouple above‐ground and below‐ground trait responses to climate change.