High exposure of global tree diversity to human pressure.

Safeguarding Earth's tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species' range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting high-priority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species' range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Best practices, errors, and perspectives of half a century of plant translocation in Italy.

Conservation translocations are becoming common conservation practice, so there is an increasing need to understand the drivers of plant translocation performance through reviews of cases at global and regional levels. The establishment of the Italian Database of Plant Translocation (IDPlanT) provides the opportunity to review the techniques used in 186 plant translocation cases performed in the last 50 years in the heart of the Mediterranean Biodiversity Hotspot. We described techniques and information available in IDPlanT and used these data to identify drivers of translocation outcomes. We tested the effect of 15 variables on survival of translocated propagules as of the last monitoring date with binomial logistic mixed-effect models. Eleven variables significantly affected survival of transplants: life form, site protection, material source, number of source populations, propagation methods, propagule life stage, planting methods, habitat suitability assessment, site preparation, aftercare, and costs. The integration of vegetation studies in the selection of suitable planting sites significantly increased the success of translocation efforts. Although posttranslocation watering had a generally positive effect on translocation outcome, other aftercare techniques did not always increase transplant survival. Finally, we found that how funds were spent appeared to be more important than the actual amount spent. Plant translocations in Italy and in the Mediterranean area should account for the complexity of speciation, gene flow, and plant migrations that has led to local adaptations and has important implications for the choice and constitution of source material.

Mejores prácticas, errores y perspectivas tras medio siglo de reubicaciones botánicas en Italia Resumen Las reubicaciones son una práctica cada vez más común en la conservación, por lo que hay una necesidad creciente por entender los factores del desempeño de las reubicaciones botánicas por medio de la revisión de casos regionales y globales. La creación de la Italian Database of Plant Translocation (IDPlanT) proporciona una oportunidad para revisar las técnicas usadas para los casos de reubicación de 186 plantas realizados durante los últimos cincuenta años en el centro del punto caliente de biodiversidad mediterránea. Describimos las técnicas y la información disponible en IDPlanT y usamos estos datos para identificar los factores involucrados en los resultados de las reubicaciones. Usamos modelos logísticos binomiales de efectos mixtos para analizar el efecto de 15 variables sobre la supervivencia de los propágulos reubicados a partir de la última fecha de monitoreo. Once de las variables afectaron de manera significativa la supervivencia de las plantas: forma de vida, protección del sitio, fuente de materiales, cantidad de poblaciones originarias, método de propagación, etapa de vida del propágulo, método de siembra, evaluación de idoneidad del hábitat, preparación del sitio, cuidados posteriores y costos. La integración de los estudios botánicos a la selección de los sitios idóneos para plantar aumentó el éxito de los esfuerzos de reubicación. Aunque el riego posterior a la reubicación tuvo un efecto positivo general sobre el resultado, las otras técnicas de cuidado posterior no siempre incrementaron la supervivencia de la planta reubicada. Por último, descubrimos que parece ser más importante cómo se utilizan los fondos que la cantidad actual empleada. Las reubicaciones botánicas en Italia y en el área del Mediterráneo deben considerar lo complejo de la especiación, el flujo génico y la migración botánica que han derivado en adaptaciones locales y que han tenido implicaciones importantes para la elección y constitución del material de origen.

Small and slow is safe: On the drought tolerance of tropical tree species.

Understanding how evolutionary history and the coordination between trait trade-off axes shape the drought tolerance of trees is crucial to predict forest dynamics under climate change. Here, we compiled traits related to drought tolerance and the fast-slow and stature-recruitment trade-off axes in 601 tropical woody species to explore their covariations and phylogenetic signals. We found that xylem resistance to embolism (P50) determines the risk of hydraulic failure, while the functional significance of leaf turgor loss point (TLP) relies on its coordination with water use strategies. P50 and TLP exhibit weak phylogenetic signals and substantial variation within genera. TLP is closely associated with the fast-slow trait axis: slow species maintain leaf functioning under higher water stress. P50 is associated with both the fast-slow and stature-recruitment trait axes: slow and small species exhibit more resistant xylem. Lower leaf phosphorus concentration is associated with more resistant xylem, which suggests a (nutrient and drought) stress-tolerance syndrome in the tropics. Overall, our results imply that (1) drought tolerance is under strong selective pressure in tropical forests, and TLP and P50 result from the repeated evolutionary adaptation of closely related taxa, and (2) drought tolerance is coordinated with the ecological strategies governing tropical forest demography. These findings provide a physiological basis to interpret the drought-induced shift toward slow-growing, smaller, denser-wooded trees observed in the tropics, with implications for forest restoration programmes.

The global spectrum of plant form and function.

Earth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today's terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Three-quarters of trait variation is captured in a two-dimensional global spectrum of plant form and function. One major dimension within this plane reflects the size of whole plants and their parts; the other represents the leaf economics spectrum, which balances leaf construction costs against growth potential. The global plant trait spectrum provides a backdrop for elucidating constraints on evolution, for functionally qualifying species and ecosystems, and for improving models that predict future vegetation based on continuous variation in plant form and function.

Climatic and evolutionary contexts are required to infer plant life history strategies from functional traits at a global scale.

Life history strategies are fundamental to the ecology and evolution of organisms and are important for understanding extinction risk and responses to global change. Using global datasets and a multiple response modelling framework we show that trait-climate interactions are associated with life history strategies for a diverse range of plant species at the global scale. Our modelling framework informs our understanding of trade-offs and positive correlations between elements of life history after accounting for environmental context and evolutionary and trait-based constraints. Interactions between plant traits and climatic context were needed to explain variation in age at maturity, distribution of mortality across the lifespan and generation times of species. Mean age at maturity and the distribution of mortality across plants' lifespan were under evolutionary constraints. These findings provide empirical support for the theoretical expectation that climatic context is key to understanding trait to life history relationships globally.

Allometric co-variation of xylem and stomata across diverse woody seedlings.

Leaf stomatal density is known to co-vary with leaf vein density. However, the functional underpinning of this relation, and how it scales to whole-plant water transport anatomy, is still unresolved. We hypothesized that the balance of water exchange between the vapour phase (in stomata) and liquid phase (in vessels) depends on the consistent scaling between the summed stomatal areas and xylem cross-sectional areas, both at the whole-plant and single-leaf level. This predicted size co-variation should be driven by the co-variation of numbers of stomata and terminal vessels. We examined the relationships of stomatal traits and xylem anatomical traits from the entire plant to individual leaves across seedlings of 53 European woody angiosperm species. There was strong and convergent scaling between total stomatal area and stem xylem area per plant and between leaf total stomatal area and midvein xylem area per leaf across all the species, irrespective of variation in leaf habit, growth-form or relative growth rate. Moreover, strong scaling was found between stomatal number and terminal vessel number, whereas not in their respective average areas. Our findings have broad implications for integrating xylem architecture and stomatal distribution and deepen our understanding of the design rules of plants' water transport network.

Assessing the reliability of predicted plant trait distributions at the global scale.

AIM: Predictions of plant traits over space and time are increasingly used to improve our understanding of plant community responses to global environmental change. A necessary step forward is to assess the reliability of global trait predictions. In this study, we predict community mean plant traits at the global scale and present a systematic evaluation of their reliability in terms of the accuracy of the models, ecological realism and various sources of uncertainty. LOCATION: Global. TIME PERIOD: Present. MAJOR TAXA STUDIED: Vascular plants. METHODS: We predicted global distributions of community mean specific leaf area, leaf nitrogen concentration, plant height and wood density with an ensemble modelling approach based on georeferenced, locally measured trait data representative of the plant community. We assessed the predictive performance of the models, the plausibility of predicted trait combinations, the influence of data quality, and the uncertainty across geographical space attributed to spatial extrapolation and diverging model predictions. RESULTS: Ensemble predictions of community mean plant height, specific leaf area and wood density resulted in ecologically plausible trait-environment relationships and trait-trait combinations. Leaf nitrogen concentration, however, could not be predicted reliably. The ensemble approach was better at predicting community trait means than any of the individual modelling techniques, which varied greatly in predictive performance and led to divergent predictions, mostly in African deserts and the Arctic, where predictions were also extrapolated. High data quality (i.e., including intraspecific variability and a representative species sample) increased model performance by 28%. MAIN CONCLUSIONS: Plant community traits can be predicted reliably at the global scale when using an ensemble approach and high-quality data for traits that mostly respond to large-scale environmental factors. We recommend applying ensemble forecasting to account for model uncertainty, using representative trait data, and more routinely assessing the reliability of trait predictions.

Mapping local and global variability in plant trait distributions.

Our ability to understand and predict the response of ecosystems to a changing environment depends on quantifying vegetation functional diversity. However, representing this diversity at the global scale is challenging. Typically, in Earth system models, characterization of plant diversity has been limited to grouping related species into plant functional types (PFTs), with all trait variation in a PFT collapsed into a single mean value that is applied globally. Using the largest global plant trait database and state of the art Bayesian modeling, we created fine-grained global maps of plant trait distributions that can be applied to Earth system models. Focusing on a set of plant traits closely coupled to photosynthesis and foliar respiration-specific leaf area (SLA) and dry mass-based concentrations of leaf nitrogen ([Formula: see text]) and phosphorus ([Formula: see text]), we characterize how traits vary within and among over 50,000 [Formula: see text]-km cells across the entire vegetated land surface. We do this in several ways-without defining the PFT of each grid cell and using 4 or 14 PFTs; each model's predictions are evaluated against out-of-sample data. This endeavor advances prior trait mapping by generating global maps that preserve variability across scales by using modern Bayesian spatial statistical modeling in combination with a database over three times larger than that in previous analyses. Our maps reveal that the most diverse grid cells possess trait variability close to the range of global PFT means.

A new method for quantifying the phylogenetic redundancy of biological communities.

The increasing use of phylogenetic methods in community ecology recognizes that accumulated evolutionary differences among species mirror, to some extent, ecological processes. The scope of this work is thus to propose a new method for the measurement of community-level phylogenetic redundancy, which takes into account the branching pattern of the underlying phylogeny. Like for functional redundancy, a measure of phylogenetic redundancy can be described as a normalized measure in the range (0-1) that relates the observed community-level phylogenetic diversity to the value of a hypothetical assemblage with the same abundance distribution of the focal community in which all species had independent evolution. Therefore, phylogenetic redundancy can be interpreted as the diversity decrease that is obtained by taking into account the evolutionary relationships among species in the calculation of diversity. The behavior of the proposed method, for which we provide a simple R function called 'phyloredundancy', was evaluated with published data on Alpine plant communities along a primary succession on a glacier foreland in northern Italy. As shown by our results, the method accounts for the length of shared branches in the phylogeny, producing a coherent framework for describing the evolutionary relationships within a species assemblage. Being based on classical diversity measures, which have been used in ecology for decades, it also has a great potential for future research in phylogenetic community ecology.

Estimation of polycyclic aromatic hydrocarbon variability in air using high volume, film, and vegetation as samplers.

Organic films and leaves provide a medium into which organic contaminants, such as polycyclic aromatic hydrocarbons (PAHs), can accumulate, resulting in a useful passive air sampler. In the present work, the temporal variability (weekly) in PAH concentrations and the fingerprint of films developed on window surfaces were investigated. Moreover, films and leaves of two tree species (Acer pseudoplatanus and Cornus mas) collected at the same time were used to derive PAH air concentrations and investigate their short-term variability. In general, the most abundant chemicals found in films were phenanthrene and pyrene (22%), followed by perylene (21%) and fluoranthene (16%), but the fingerprint (in contrast to leaves and air) changed over time. Leaf derived air concentrations were within a factor of 2 to 9 from measured values, while air concentrations back-calculated from films were within a factor of 2 to 53. This happened because predicted air concentrations using films and vegetation samplers (especially for low KOA chemicals) generally reflect only the last few hours (due to the fast equilibrium) of the weekly integrated samples obtained employing the high-volume sampler. This means that films and leaves can be usefully employed for predicting the short-term variability of low KOA organic contaminant air concentrations.

Using Plant Functional Traits to Define the Biomass Energy Potential of Invasive Alien Plant Species.

The eradication of invasive alien plant species (IAPS) is mandatory worldwide, but the resulting biomass is still considered waste. The energy use of biomasses obtained from IAPS eradication may represent ecological and economic benefits, creating synergies with restoration projects. We evaluated whether the growth forms and functional types identified using the functional space of 63 IAPS corresponded to a possible bioenergy use through multivariate analysis techniques. We extracted leaf and nutrient traits and Grime's CSR plant strategies from an existing database. We calculated the carbon-to-nitrogen ratio (C:N) and gross heating value (GHV) as indicators of biochemical or thermal processes, respectively. For 10 species, we measured the above-ground biomass C:N and GHV (including leaves, stems and branches) and correlated them with those of leaves and with plant adaptive strategies. We identified four groups of IAPS indicative of the main trade-offs between plant economics and size variation, which respectively correlated with C:N and GHV. Herbaceous IAPS were better suited to biochemical processes, and woody IAPS to thermal ones. Overall, Grime's CSR strategies were the best tool to define the IAPS bioenergy potential. In the long term, competitive and ruderal IAPSs can represent a reusable feedstock until their complete eradication.

Plant traits poorly predict winner and loser shrub species in a warming tundra biome.

Climate change is leading to species redistributions. In the tundra biome, shrubs are generally expanding, but not all tundra shrub species will benefit from warming. Winner and loser species, and the characteristics that may determine success or failure, have not yet been fully identified. Here, we investigate whether past abundance changes, current range sizes and projected range shifts derived from species distribution models are related to plant trait values and intraspecific trait variation. We combined 17,921 trait records with observed past and modelled future distributions from 62 tundra shrub species across three continents. We found that species with greater variation in seed mass and specific leaf area had larger projected range shifts, and projected winner species had greater seed mass values. However, trait values and variation were not consistently related to current and projected ranges, nor to past abundance change. Overall, our findings indicate that abundance change and range shifts will not lead to directional modifications in shrub trait composition, since winner and loser species share relatively similar trait spaces.

Global beta-diversity of angiosperm trees is shaped by Quaternary climate change.

As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleoclimate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (richness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta-diversity in regions that experienced large temperature change, reflecting phylogenetically and functionally selective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide.

Combined use of leaf size and economics traits allows direct comparison of hydrophyte and terrestrial herbaceous adaptive strategies.

BACKGROUND AND AIMS: Hydrophytes generally exhibit highly acquisitive leaf economics. However, a range of growth forms is evident, from small, free-floating and rapidly growing Lemniden to large, broad-leaved Nymphaeiden, denoting variability in adaptive strategies. Traits used to classify adaptive strategies in terrestrial species, such as canopy height, are not applicable to hydrophytes. We hypothesize that hydrophyte leaf size traits and economics exhibit sufficient overlap with terrestrial species to allow a common classification of plant functional types, sensu Grime's CSR theory. METHODS: Leaf morpho-functional traits were measured for 61 species from 47 water bodies in lowland continental, sub-alpine and alpine bioclimatic zones in southern Europe and compared against the full leaf economics spectrum and leaf size range of terrestrial herbs, and between hydrophyte growth forms. KEY RESULTS: Hydrophytes differed in the ranges and mean values of traits compared with herbs, but principal components analysis (PCA) demonstrated that both groups shared axes of trait variability: PCA1 encompassed size variation (area and mass), and PCA2 ranged from relatively dense, carbon-rich leaves to nitrogen-rich leaves of high specific leaf area (SLA). Most growth forms exhibited trait syndromes directly equivalent to herbs classified as R adapted, although Nymphaeiden ranged between C and SR adaptation. CONCLUSIONS: Our findings support the hypothesis that hydrophyte adaptive strategy variation reflects fundamental trade-offs in economics and size that govern all plants, and that hydrophyte adaptive strategies can be directly compared with terrestrial species by combining leaf economics and size traits.

Mulching in lowland hay meadows drives an adaptive convergence of above- and below-ground traits reducing plasticity and improving biomass: A possible tool for enhancing phytoremediation.

We aimed to understand the effect of mulching (i.e., cutting and leaving the crushed biomass to decompose in situ) on above- and below-ground plant functional traits and whether this practice may be a potential tool for enhancing the phytoremediation of lowland hay meadows. To this aim, we evaluated at the community level seven years of mulching application in a PCBs and HMs soil-polluted Site of National Interest (SIN Brescia-Caffaro) through the analysis of the floristic composition and the above- and below-ground plant traits. We found that the abandonment of agricultural activities led to a marked increase in the soil organic carbon and pH, and the over-imposed mulching additionally induced a slight increase in soil nutrients. Mulching favored the establishment of a productive plant community characterized by a more conservative-resource strategy, a higher biomass development, and lower plasticity through an adaptative convergence between above- and below-ground organs. In particular, the analysis of the root depth distribution highlighted the key role of roots living in the upper soil layer (10 cm). Mulching did not show a significant effect on plant species known to be effective in terms of PCB phytoremediation. However, the mulching application appears to be a promising tool for enhancing the root web that functions as the backbone for the proliferation of microbes devoted to organic contaminants' degradation and selects a two-fold number of plant species known to be metal-tolerant. However, besides these potential positive effects of the mulching application, favoring species with a higher biomass development, in the long term, may lead to a biodiversity reduction and thus to potential consequences also on the diversity of native species important for the phytoremediation.

Global relationships in tree functional traits.

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide.

The global spectrum of plant form and function: enhanced species-level trait dataset.

Here we provide the 'Global Spectrum of Plant Form and Function Dataset', containing species mean values for six vascular plant traits. Together, these traits -plant height, stem specific density, leaf area, leaf mass per area, leaf nitrogen content per dry mass, and diaspore (seed or spore) mass - define the primary axes of variation in plant form and function. The dataset is based on ca. 1 million trait records received via the TRY database (representing ca. 2,500 original publications) and additional unpublished data. It provides 92,159 species mean values for the six traits, covering 46,047 species. The data are complemented by higher-level taxonomic classification and six categorical traits (woodiness, growth form, succulence, adaptation to terrestrial or aquatic habitats, nutrition type and leaf type). Data quality management is based on a probabilistic approach combined with comprehensive validation against expert knowledge and external information. Intense data acquisition and thorough quality control produced the largest and, to our knowledge, most accurate compilation of empirically observed vascular plant species mean traits to date.

Climatic and soil factors explain the two-dimensional spectrum of global plant trait variation.

Plant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land-climate feedbacks. Still, we lack a global understanding of how land and climate affect plant traits. A previous global analysis of six traits observed two main axes of variation: (1) size variation at the organ and plant level and (2) leaf economics balancing leaf persistence against plant growth potential. The orthogonality of these two axes suggests they are differently influenced by environmental drivers. We find that these axes persist in a global dataset of 17 traits across more than 20,000 species. We find a dominant joint effect of climate and soil on trait variation. Additional independent climate effects are also observed across most traits, whereas independent soil effects are almost exclusively observed for economics traits. Variation in size traits correlates well with a latitudinal gradient related to water or energy limitation. In contrast, variation in economics traits is better explained by interactions of climate with soil fertility. These findings have the potential to improve our understanding of biodiversity patterns and our predictions of climate change impacts on biogeochemical cycles.

The association of leaf sulfur content with the leaf economics spectrum and plant adaptive strategies.

Sulfur is an essential macronutrient for plant primary metabolism. Its availability can modulate plant growth in most terrestrial ecosystems. However, its relationship with other leaf and nutrient traits, and hence its contribution to plant functioning, remains unclear. We analysed leaf and nutrient traits for 740 vascular plant species growing in a wide range of environmental conditions in Northern Italy. We determined whether leaf sulfur content per unit leaf dry mass (LSC) is associated with leaf economics spectrum, and whether its distribution among functional types (growth forms, leaf life span categories, and Grime's CSR (Competitive, Stress-tolerant, Ruderal strategies) could help to elucidate adaptive differences within plant taxa. High LSC values were mainly associated with fast-growing species representative of R- and C- strategy selection, thus the acquisitive extreme of plant economics, reflecting strong potential connections with ecosystem properties such as biomass production or litter decomposability. In general, LSC was significantly and positively correlated with leaf nitrogen content, and nitrogen to sulfur ratio was constant throughout growth forms, leaf life span and CSR strategies, and phylogenetic effects were evident. Our findings highlight that LSC variation is strongly associated with the leaf economics spectrum, suggesting that additional nutrients seldom included in functional analyses may also be embroiled within the context of plant economics. However, different ratios among nitrogen and sulfur may be expected across different plant families, suggesting that deeper insight from functional groups can provide a bridge between plant stoichiometry and ecology, useful for the evaluation of ecological responses to global change.