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The effects of climate change on tropical forests will depend on how diverse tropical tree species respond to drought. Current distributions of evergreen and deciduous tree species across local and regional moisture gradients reflect their ability to tolerate drought stress, and might be explained by functional traits. We measured leaf water potential at turgor loss (i.e. ‘wilting point’; πtlp), wood density (WD) and leaf mass per area (LMA) on 50 of the most abundant tree species in central Panama. We then tested their ability to explain distributions of evergreen and deciduous species within a 50 ha plot on Barro Colorado Island and across a 70 km rainfall gradient spanning the Isthmus of Panama. Among evergreen trees, species with lower πtlp were associated with drier habitats, with πtlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, πtlp did not predict habitat associations among deciduous species. Across spatial scales, πtlp is a useful indicator of habitat preference for tropical tree species that retain their leaves during periods of water stress, and holds the potential to predict vegetation responses to climate change.

Understanding how tropical tree phenology (i.e., the timing and amount of seed and leaf production) responds to climate is vital for predicting how climate change may alter ecological functioning of tropical forests. We examined the effects of temperature, rainfall, and photosynthetically active radiation (PAR) on seed phenology of four dominant species and community‐level leaf phenology in a montane wet forest on the island of Hawaiʻi using monthly data collected over ~ 6 years. We expected that species phenologies would be better explained by variation in temperature and PAR than rainfall because rainfall at this site is not limiting. The best‐fit model for all four species included temperature, rainfall, and PAR. For three species, including two foundational species of Hawaiian forests (Acacia koa and Metrosideros polymorpha), seed production declined with increasing maximum temperatures and increased with rainfall. Relationships with PAR were the most variable across all four species. Community‐level leaf litterfall decreased with minimum temperatures, increased with rainfall, and showed a peak at PAR of ~ 400 μmol/m2s−1. There was considerable variation in monthly seed and leaf production not explained by climatic factors, and there was some evidence for a mediating effect of daylength. Thus, the impact of future climate change on this forest will depend on how climate change interacts with other factors such as daylength, biotic, and/or evolutionary constraints. Our results nonetheless provide insight into how climate change may affect different species in unique ways with potential consequences for shifts in species distributions and community composition.

In urban Mediterranean ecosystems, invasive alien plants should have characteristics that enable faster resource acquisition and utilization than native species. We tested this hypothesis by comparing stem and leaf functional traits from five abundant native woody shrubs and five of the most abundant alien species within an urban coastal scrub community in Ensenada, Baja California, Mexico. Twelve anatomical and ecophysiological traits were studied: light-saturated CO2 assimilation rate, stomatal conductance, stem and leaf water potentials during transpiration, % stem allocation to bark, xylem and pith, stem xylem vessel diameter and density, leaf size, and % allocation to spongy and palisade mesophyll. Alien species varied more in these traits, and their trait combinations reflected less investment in long-lived tissues; they had higher CO2 assimilation and stomatal conductance rates, higher values of leaf and stem water potential, and higher allocation to pith, larger stem xylem vessels, and larger leaf areas. In contrast, native woody shrub species were relatively convergent in leaf and stem traits reflecting more conservative resource use. Within disturbed urban environments, alien species may outcompete the native stress-tolerant species through rapid resource acquisition, enhanced by a broader set of functional trait combinations across species.

We review possible physiological, anatomical, and evolutionary limitations that may explain the rarity of C4 photosynthesis in trees and discuss how the C4 trees in Euphorbia are exceptions to this., Since C4 photosynthesis was first discovered >50 years ago, researchers have sought to understand how this complex trait evolved from the ancestral C3 photosynthetic machinery on >60 occasions. Despite its repeated emergence across the plant kingdom, C4 photosynthesis is notably rare in trees, with true C4 trees only existing in Euphorbia. Here we consider aspects of the C4 trait that could limit but not preclude the evolution of a C4 tree, including reduced quantum yield, increased energetic demand, reduced adaptive plasticity, evolutionary constraints, and a new theory that the passive symplastic phloem loading mechanism observed in trees, combined with difficulties in maintaining sugar and water transport over a long pathlength, could make C4 photosynthesis largely incompatible with the tree lifeform. We conclude that the transition to a tree habit within C4 lineages as well as the emergence of C4 photosynthesis within pre-existing trees would both face a series of challenges that together explain the global rarity of C4 photosynthesis in trees. The C4 trees in Euphorbia are therefore exceptional in how they have circumvented every potential barrier to the rare C4 tree lifeform.

Drought is a major stress for plants, creating a strong selection pressure for traits that enable plant growth and survival in dry environments. Many drought responses are conserved species-wide responses, while others vary among populations distributed across heterogeneous environments. We tested how six populations of the widely distributed California valley oak (Quercus lobata) sampled from contrasting climates would differ in their response to soil drying relative to well-watered controls in a common environment by measuring ecophysiological traits in 93 individuals and gene expression (RNA-seq) in 42 individuals. Populations did not differ in their adjustment of turgor loss point during soil drying, suggesting a generalized species-wide response. Differential expression analysis identified 689 genes with a common response to treatment across populations and 470 genes with population-specific responses. Weighted gene co-expression network analysis (WGCNA) identified groups of genes with similar expression patterns that may be regulated together (gene modules). Several gene modules responded differently to water stress among populations, suggesting regional differences in gene network regulation. Populations from sites with a high mean annual temperature responded to the imposed water stress with significantly greater changes in gene module expression, indicating that these populations may be locally adapted to respond to drought. We propose that this variation among valley oak populations provides a mechanism for differential tolerance to the increasingly frequent and severe droughts in California.

The maximum stomatal conductance (g), a major anatomical constraint on plant productivity, is a function of the stomatal area fraction (f) and stomatal space-use efficiency (e). However, f and g have been considered as equivalents, with e rarely considered, and their adaptation to the environment and their regulation of ecosystem productivity are unclear. Here, we analyzed the community-weighted mean, variance, skewness, and kurtosis of stomatal traits from tropical to cold-temperature forests. The variance of g and f was higher for arid sites, indicating greater functional niche differentiation, whereas that for e was lower, indicating convergence in efficiency. Besides, when other stomatal trait distributions remained unchanged, increasing kurtosis but decreasing skewness of g would improve ecosystem productivity, and f showed the opposite patterns. These findings highlight how the relative importance and equivalence of inter-related traits can differ at community scale.

Droughts in a warming climate have become more common and more extreme, making understanding forest responses to water stress increasingly pressing. Analysis of water stress in trees has long focused on water potential in xylem and leaves, which influences stomatal closure and water flow through the soil-plant-atmosphere continuum. At the same time, changes of vegetation water content (VWC) are linked to a range of tree responses, including fluxes of water and carbon, mortality, flammability, and more. Unlike water potential, which requires demanding in situ measurements, VWC can be retrieved from remote sensing measurements, particularly at microwave frequencies using radar and radiometry. Here, we highlight key frontiers through which VWC has the potential to significantly increase our understanding of forest responses to water stress. To validate remote sensing observations of VWC at landscape scale and to better relate them to data assimilation model parameters, we introduce an ecosystem-scale analog of the pressure–volume curve, the non-linear relationship between average leaf or branch water potential and water content commonly used in plant hydraulics. The sources of variability in these ecosystem-scale pressure-volume curves and their relationship to forest response to water stress are discussed. We further show to what extent diel, seasonal, and decadal dynamics of VWC reflect variations in different processes relating the tree response to water stress. VWC can also be used for inferring below-ground conditions—which are difficult to impossible to observe directly. Lastly, we discuss how a dedicated geostationary spaceborne observational system for VWC, when combined with existing datasets, can capture diel and seasonal water dynamics to advance the science and applications of global forest vulnerability to future droughts.

Some lineages radiate spectacularly when colonizing a region, but others do not. Large radiations are often attributed to species’ adaptation into niches, or to other drivers, such as biogeography including dispersal ability and spatial structure of the landscape. Here we aim to disentangle the factors determining radiation size, by modeling simplified scenarios without the complexity of explicit niches. We build a spatially structured neutral model free from niches and incorporating a form of protracted speciation that accounts for gene flow between populations. We find that a wide range of radiation sizes are possible in this model depending on the combination of geographic isolation and species’ dispersal ability. At extremely low rates of dispersal between patches, each patch maintains its own endemic species. Intermediate dispersal rates foster larger radiations as they allow occasional movement between patches whilst sufficiently restricting gene flow to support further speciation in allopatry. As dispersal rates increase further, a critical point is reached at which demographically identical lineages may vary greatly in radiation size due to rare and stochastic dispersal events. At the critical point in dispersal frequency, some lineages remain a single species for a comparatively long time, whilst others with identical characteristics produce the largest radiations of all via a new mechanism for rapid radiation that we term a ‘radiation cascade’. Given a single species covering many patches connected with gene flow, a radiation cascade is triggered when stochastic dispersal is unusually low for a period, leading to an initial speciation event. This speciation means there are fewer individuals per species and thus further reduced gene flow between conspecifics. Reduced gene flow in turn makes it easier for further speciation to occur. During a radiation cascade, dispersal of individuals between patches continues at the same rate as before, but due to the increasing diversity it primarily introduces novel species that will later speciate, rather than adding to gene flow of existing species. Once a radiation cascade begins, it continues rapidly until it is arrested by a new equilibrium between speciation and extinction. We speculate that such radiation cascades may occur more generally and are not only present in neutral models. This process may help to explain rapid radiation, and the extreme radiation sizes of certain lineages with dispersing ancestors. Whilst niches no doubt play a role in community assembly, our findings lead us to question whether diversification and adaptation into niches is sometimes an effect of speciation and rapid radiation, rather than its cause.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur.

Symbiotic nitrogen (N)-fixing trees can provide large quantities of new N to ecosystems, but only if they are sufficiently abundant. The overall abundance and latitudinal abundance distributions of N-fixing trees are well characterised in the Americas, but less well outside the Americas. Here, we characterised the abundance of N-fixing trees in a network of forest plots spanning five continents, ~5,000 tree species and ~4 million trees. The majority of the plots (86%) were in America or Asia. In addition, we examined whether the observed pattern of abundance of N-fixing trees was correlated with mean annual temperature and precipitation. Outside the tropics, N-fixing trees were consistently rare in the forest plots we examined. Within the tropics, N-fixing trees were abundant in American but not Asian forest plots (~7% versus ~1% of basal area and stems). This disparity was not explained by mean annual temperature or precipitation. Our finding of low N-fixing tree abundance in the Asian tropics casts some doubt on recent high estimates of N fixation rates in this region, which do not account for disparities in N-fixing tree abundance between the Asian and American tropics. Synthesis. Inputs of nitrogen to forests depend on symbiotic nitrogen fixation, which is constrained by the abundance of N-fixing trees. By analysing a large dataset of ~4 million trees, we found that N-fixing trees were consistently rare in the Asian tropics as well as across higher latitudes in Asia, America and Europe. The rarity of N-fixing trees in the Asian tropics compared with the American tropics might stem from lower intrinsic N limitation in Asian tropical forests, although direct support for any mechanism is lacking. The paucity of N-fixing trees throughout Asian forests suggests that N inputs to the Asian tropics might be lower than previously thought.

Leaf density links the resistance to mechanical and hydraulic stress in Mediterranean shrubs as it is associated with the water potential at turgor loss and the moduli of elasticity and strength. Understanding the patterns of hydraulic and mechanical trait variation in vascular plants is critical to predicting species’ stress tolerance. Although previous work has shown that hydraulic and mechanical traits are decoupled in stems, there is little information available for leaves, which are organs more diversified in structure, function, and possibly drought tolerance strategies across habitats. We tested for coordination between leaf hydraulic traits related to drought tolerance and the mechanical resistance of leaves, for 17 shrub species from the arid and semiarid vegetation of the California Floristic Province. Bayesian and phylogenetic correlations showed that across species, hydraulic and mechanical traits both had strong associations with the water potential at turgor loss, and with leaf tissue density. However, leaf maximum hydraulic conductance and the water potential at 50% and 80% loss of hydraulic conductance were statistically independent of two key mechanical traits, the leaf modulus of elasticity and leaf structural strength. Our results suggest that leaf biomechanical traits, which reflect construction costs and contribute to leaf longevity, are decoupled from hydraulic capacity and safety. The independence of hydraulic and mechanical protection in leaves enables a wide range of trait combinations in leaves, which would increase their adaptive potential across ecosystems.

[Motivation]: Trait data are fundamental to the quantitative description of plant form and function. Although root traits capture key dimensions related to plant responses to changing environmental conditions and effects on ecosystem processes, they have rarely been included in large-scale comparative studies and global models. For instance, root traits remain absent from nearly all studies that define the global spectrum of plant form and function. Thus, to overcome conceptual and methodological roadblocks preventing a widespread integration of root trait data into large-scale analyses we created the Global Root Trait (GRooT) Database. GRooT provides ready to use data by combining the expertise of root ecologists with data mobilization and curation. Specifically, we (a) determined a set of core root traits relevant to the description of plant form and function based on an assessment by experts, (b) maximized species coverage through data standardization within and among traits, and (c) implemented data quality checks., [Main types of variables contained]: GRooT contains 114,222 trait records on 38 continuous root traits.Spatial location and grain: Global coverage with data from arid, continental, polar, temperate and tropical biomes. Data on root traits were derived from experimental studies and field studies., [Time period and grain]: Data were recorded between 1911 and 2019., [Major taxa and level of measurement]: GRooT includes root trait data for which taxonomic information is available. Trait records vary in their taxonomic resolution, with subspecies or varieties being the highest and genera the lowest taxonomic resolution available. It contains information for 184 subspecies or varieties, 6,214 species, 1,967 genera and 254 families. Owing to variation in data sources, trait records in the database include both individual observations and mean values., [Software format]: GRooT includes two csv files. A GitHub repository contains the csv files and a script in R to query the database., The study has been supported by the TRY initiative on plant traits (http://www.try-db.org). The TRY initiative and database are hosted, developed and maintained at the Max Planck Institute for Biogeochemistry, Jena, Germany. TRY is currently supported by DIVERSITAS/Future Earth and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig. Open access funding enabled and organized by Projekt DEAL.

Increases in the frequency and severity of droughts across many regions worldwide necessitate an improved capacity to determine the water status of plants at organ, whole plant, canopy, and regional scales. Noninvasive methods have most potential for simultaneously improving basic water relations research and ground‐, flight‐, and space‐based sensing of water status, with applications in sustainability, food security, and conservation. The most frequently used methods to measure the most salient proxies of plant water status, that is, water mass per leaf area (WMA), relative water content (RWC), and leaf water potential (Ψleaf), require the excision of tissues and laboratory analysis, and have thus been limited to relatively low throughput and small study scales. Applications using electromagnetic radiation in the visible, infrared, and terahertz ranges can resolve the water status of canopies, yet heretofore have typically focused on statistical approaches to estimating RWC for leaves before and after severe dehydration, and few have predicted Ψleaf. Terahertz radiation has great promise to estimate leaf water status across the range of leaf dehydration important for the control of gas exchange and leaf survival. We demonstrate a refined method and physical model to predict WMA, RWC, and Ψleaf from terahertz transmission across a wide range of levels of dehydration for given leaves of three species, as well as across leaves of given species and across multiple species. These findings highlight the powerful potential and the outstanding challenges in applying in vivo terahertz spectrometry as a remote sensor of water status for a range of applications.

Plants often experience multiple stresses in a given day or season, and it is self-evident that given functional traits can provide tolerances of multiple stresses. Yet, the multiple functions of individual traits are rarely explicitly considered in ecology and evolution due to a lack of a quantitative framework. We present a theory for considering the combined importance of the several functions that a single trait can contribute to alleviating multiple stresses. We derive five inter-related general predictions: (1) that trait multifunctionality is overall highly beneficial to fitness; (2) that species possessing multifunctional traits should increase in abundance and in niche breadth; (3) that traits are typically optimized for multiple functions and thus can be far from optimal for individual functions; (4) that the relative importance of each function of a multifunctional trait depends on the environment; and (5) that traits will be often “co-opted” for additional functions during evolution and community assembly. We demonstrate how the theory can be applied quantitatively by examining the multiple functions of leaf trichomes (hairs) using heuristic model simulations, substantiating the general principles. We identify avenues for further development and applications of the theory of trait multifunctionality in ecology and evolution.

© 2018, The Author(s). Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four ‘survival modes’ that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Leaf chlorophyll is essential to harvest light energy to drive photosynthesis. Yet, most of studies of leaf chlorophyll concentration (Chl) have typically been limited to few species or sites, and there has been little understanding of its variation among species of contrasting environments, or across forests differing in gross primary productivity (GPP). We measured mass-based foliar Chl concentrations (Chl a, Chl b, and total Chl) for 937 species of nine Chinese forests ranging from tropical to cold-temperate regions. Total Chl varied by over ten-fold from 1.45 to 19.2 mg g−1 (mean: 5.54 mg g−1), and differed among plant functional groups. At the community level, total Chl decreased with increasing latitude. The ratio of Chl a/b decreased from trees to shrubs to herbs, consistent with adaptation to contrasting irradiance, i.e., overstory vs. understory. Total Chl was positively related to leaf N and P concentrations, as would be expected to optimize photosynthesis and carbon fixation. Across forests, GPP was positively correlated with community-averaged Chl (r = 0.57; P

1.Stomata control carbon and water vapor exchange between leaves and the atmosphere, thus it can influence water use efficiency and reflect plant adaptation to climate. However, the spatial patterns of leaf stomatal traits and relationships between stomatal trait and water use efficiency across natural communities remain unclear. 2.We measured stomatal density, stomatal size, and stomatal area fraction for 737 plant species from nine forests ranging from tropical to cold-temperate forests. 3.Stomatal density, stomatal size, and stomatal area fraction were all log-normally distributed, and different across species, plant functional groups (trees, shrubs, and herbs), and communities. At the regional scale, variation in stomatal traits was primarily related to species, followed by climate and soil types. 4.The community-weighted mean of stomatal size increased linearly with latitude, whereas those of stomatal density and stomatal area fraction showed humpbacked relationship. The community-weighted mean of stomatal area fraction was correlated with climatic aridity, consistent with the adaptation strategies of plant species to achieve high maximum rates of gas exchange in arid regions when water is available. Further, community-weighted mean of stomatal area fraction was positively correlated with water use efficiency in natural forest communities, indicating that plants have lower stomatal conductance in order to adapt greater aridity conditions. 5.These findings highlight the strong associations of stomatal traits with plant functional group and climate at a regional scale, representing the adaptation strategies of stomatal traits across natural forest communities to climate. This article is protected by copyright. All rights reserved.

Leaf size varies by over a 100,000-fold among species worldwide. Although 19th-century plant geographers noted that the wet tropics harbor plants with exceptionally large leaves,the latitudinal gradient of leaf size has not been well quantified nor the key climatic drivers convincingly identified. Here, we characterize worldwide patterns in leaf size. Large-leaved species predominate in wet, hot, sunny environments; small-leaved species typify hot,sunny environments only in arid conditions; small leaves are also found in high latitudes and elevations. By modeling the balance of leaf energy inputs and outputs, we show that daytime and nighttime leaf-to-air temperature differences are key to geographic gradients in leaf size. This knowledge can enrich ?next-generation? vegetation models in which leaf temperature and water use during photosynthesis play key roles. Fil: Wright, Ian J.. Macquarie University; Australia Fil: Dong, Ning. Macquarie University; Australia. University of Reading, Whiteknights; Reino Unido Fil: Maire, Vincent. Université du Québec à Trois-Rivières; Canadá. Macquarie University; Australia Fil: Prentice, I. Colin. Macquarie University; Australia. Imperial College London; Reino Unido Fil: Westoby, Mark. Macquarie University; Australia Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Gallagher, Rachel V.. Macquarie University; Australia Fil: Jacobs, Bonnie F.. Southern Methodist University; Estados Unidos Fil: Kooyman, Robert. Macquarie University; Australia Fil: Law, Elizabeth A.. Macquarie University; Australia. The University Of Queensland; Australia Fil: Leishman, Michelle R.. Macquarie University; Australia Fil: Niinemets, Ülo. Estonian University of Life Sciences; Estonia Fil: Reich, Peter B.. University of Minnesota; Estados Unidos. Western Sydney University; Australia Fil: Sack, Lawren. University of California. Department of Ecology and Evolutionary Biology; Estados Unidos Fil: Villar, Rafael. Universidad de Córdoba. Facultad de Ciencias. Área de Ecología; España Fil: Wang, Han. Northwest A & F University. College of Forestry; China Fil: Wilf, Peter. Pennsylvania State University. Department of Geosciences; Estados Unidos

Maintaining tree diversity Negative interaction among plant species is known as conspecific negative density dependence (CNDD). This ecological pattern is thought to maintain higher species diversity in the tropics. LaManna et al. tested this hypothesis by comparing how tree species diversity changes with the intensity of local biotic interactions in tropical and temperate latitudes (see the Perspective by Comita). Stronger local specialized biotic interactions seem to prevent erosion of biodiversity in tropical forests, not only by limiting populations of common species, but also by strongly stabilizing populations of rare species, which tend to show higher CNDD in the tropics. Science , this issue p. 1389 ; see also p. 1328

Climate change is expected to exacerbate drought for many plants, making drought tolerance a key driver of species and ecosystem responses. Plant drought tolerance is determined by multiple traits, but the relationships among traits, either within individual plants or across species, have not been evaluated for general patterns across plant diversity. We synthesized the published data for stomatal closure, wilting, declines in hydraulic conductivity in the leaves, stems, and roots, and plant mortality for 262 woody angiosperm and 48 gymnosperm species. We evaluated the correlations among the drought tolerance traits across species, and the general sequence of water potential thresholds for these traits within individual plants. The trait correlations across species provide a framework for predicting plant responses to a wide range of water stress from one or two sampled traits, increasing the ability to rapidly characterize drought tolerance across diverse species. Analyzing these correlations also identified correlations among the leaf and stem hydraulic traits and the wilting point, or turgor loss point, beyond those expected from shared ancestry or independent associations with water stress alone. Further, on average, the angiosperm species generally exhibited a sequence of drought tolerance traits that is expected to limit severe tissue damage during drought, such as wilting and substantial stem embolism. This synthesis of the relationships among the drought tolerance traits provides crucial, empirically supported insight into representing variation in multiple traits in models of plant and ecosystem responses to drought.

Stomatal conductance (g

1. Climate change is a world-wide threat to biodiversity and ecosystem structure, functioning and services. To understand the underlying drivers and mechanisms, and to predict the consequences for nature and people, we urgently need better understanding of the direction and magnitude of climate change impacts across the soil–plant–atmosphere continuum. An increasing number of climate change studies are creating new opportunities for meaningful and high-quality generalizations and improved process understanding. However, significant challenges exist related to data availability and/or compatibility across studies, compromising opportunities for data re-use, synthesis and upscaling. Many of these challenges relate to a lack of an established ‘best practice’ for measuring key impacts and responses. This restrains our current understanding of complex processes and mechanisms in terrestrial ecosystems related to climate change. 2. To overcome these challenges, we collected best-practice methods emerging from major ecological research networks and experiments, as synthesized by 115 experts from across a wide range of scientific disciplines. Our handbook contains guidance on the selection of response variables for different purposes, protocols for standardized measurements of 66 such response variables and advice on data management. Specifically, we recommend a minimum subset of variables that should be collected in all climate change studies to allow data re-use and synthesis, and give guidance on additional variables critical for different types of synthesis and upscaling. The goal of this community effort is to facilitate awareness of the importance and broader application of standardized methods to promote data re-use, availability, compatibility and transparency. We envision improved research practices that will increase returns on investments in individual research projects, facilitate second-order research outputs and create opportunities for collaboration across scientific communities. Ultimately, this should significantly improve the quality and impact of the science, which is required to fulfil society's best practice, coordinated experiments, data management and documentation, ecosystem, experimental macroecology, methodology, open science, vegetation This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2019 The Authors. Methods in Ecology and Evolution published by John Wiley & Sons Ltd on behalf of British Ecological Society.

International audience; Plants are enormously diverse in their traits and ecological adaptation, even within given ecosystems, such as tropical rainforests. Accounting for this diversity in vegetation models poses serious challenges. Global plant functional trait databases have highlighted general trait correlations across species that have considerably advanced this research program. However, it remains unclear whether trait correlations found globally hold within communities, and whether they extend to drought tolerance traits. For 134 individual plants spanning a range of sizes and life forms (tree, liana, understory species) within an Amazonian forest, we measured leaf drought tolerance (leaf water potential at turgor loss point, πtlp), together with 17 leaf traits related to various functions, including leaf economics traits and nutrient composition (leaf mass per area, LMA; and concentrations of C, N, P, K, Ca, and Mg per leaf mass and area), leaf area, water use efficiency (carbon isotope ratio), and time-integrated stomatal conductance and carbon assimilation rate per leaf mass and area. We tested trait coordination and the ability to estimate πtlp from the other traits through model selection. Performance and transferability of the best predictive model were assessed through cross-validation. πtlp was positively correlated with leaf area, and with N, P and K concentrations per leaf mass, but not with LMA or any other studied trait. Five axes were needed to account for > 80% of trait variation, but only three of them explained more variance than expected at random. The best model explained only 30% of the variation in πtlp, and out-sample predictive performance was variable across life forms or canopy strata, suggesting a limited transferability of the model. Synthesis. We found a weak correlation among leaf drought tolerance and other leaf traits within a forest community. We conclude that higher trait dimensionality than assumed under the leaf economics spectrum may operate among leaves within plant communities, with important implications for species coexistence and responses to changing environmental conditions, and also for the representation of community diversity in vegetation models.

Drought-induced tree mortality has been observed globally and is expected to increase under climate change scenarios, with large potential consequences for the terrestrial carbon sink. Predicting mortality across species is crucial for assessing the effects of climate extremes on forest community biodiversity, composition, and carbon sequestration. However, the physiological traits associated with elevated risk of mortality in diverse ecosystems remain unknown, although these traits could greatly improve understanding and prediction of tree mortality in forests. We performed a meta-analysis on species' mortality rates across 475 species from 33 studies around the globe to assess which traits determine a species' mortality risk. We found that species-specific mortality anomalies from community mortality rate in a given drought were associated with plant hydraulic traits. Across all species, mortality was best predicted by a low hydraulic safety margin-the difference between typical minimum xylem water potential and that causing xylem dysfunction-and xylem vulnerability to embolism. Angiosperms and gymnosperms experienced roughly equal mortality risks. Our results provide broad support for the hypothesis that hydraulic traits capture key mechanisms determining tree death and highlight that physiological traits can improve vegetation model prediction of tree mortality during climate extremes.

Summary Conspecific density may contribute to patterns of species assembly through negative density dependence (NDD) as predicted by the Janzen-Connell hypothesis, or through facilitation (positive density dependence; PDD). Conspecific density effects are expected to be more negative in darker and wetter environments due to higher pathogen abundance and more positive in stressful, especially dry, environments (stress-gradient hypothesis). For NDD to contribute to maintaining diversity, it should be apparent at the community-wide scale as a negative correlation between seedling recruitment, growth or survival and conspecific adult abundance (community compensatory trend; CCT). We examined seedling survival in relation to con- and heterospecific adults within 10 m and con- and heterospecific seedlings within 1 m for 13 species within two 4-ha permanent plots located in dry and wet forests in Hawaii. We also examined interactions between conspecific density and light and species’ commonness. For all species pooled, adult conspecific effects were positive (PDD) in both dry and wet forests, though they were stronger in the dry forest. In contrast, seedling conspecific effects were negative (NDD), though only significantly so in the wet forest. The strength and direction of density effects varied with understorey light such that seedlings had the highest survival where both adult conspecific density and light were high but the lowest survival where seedling conspecific density and light were high. In the wet forest, the most common species showed positive effects of adult conspecifics, but the less common species showed negative adult conspecific effects. We found mixed evidence for a CCT: seedling survival was positively correlated with basal area, but negatively correlated with tree density (stems ha−1). Thus, it remains unclear whether NDD is a diversity-maintaining mechanism in these forests. Synthesis. Overall, we found that positive conspecific effects influenced seedling mortality patterns more than negative interactions did, even in tropical wet forest where NDD is predicted to drive species’ abundances. Additionally, the strength and direction of density effects varied with forest type, PAR, and species’ abundance, underscoring the need to consider abiotic factors and species’ life-history traits in tests of density dependence hypotheses.

Spatial patterns in trait variation reflect underlying community assembly processes, allowing us to test hypotheses about their trait and environmental drivers by identifying the strongest correlates of characteristic spatial patterns. For 43 evergreen tree species (> 1 cm dbh) in a 20-ha seasonal tropical rainforest plot in Xishuangbanna, China, we compared the ability of drought-tolerance traits, other physiological traits, and commonly measured functional traits to predict the spatial patterns expected from the assembly processes of habitat associations, niche-overlap-based competition, and hierarchical competition. We distinguished the neighborhood-scale (0-20 m) patterns expected from competition from larger-scale habitat associations with a wavelet method. Species' drought tolerance and habitat variables related to soil water supply were strong drivers of habitat associations, and drought tolerance showed a significant spatial signal for influencing competition. Overall, the traits most strongly associated with habitat, as quantified using multivariate models, were leaf density, leaf turgor loss point (π(tlp); also known as the leaf wilting point), and stem hydraulic conductivity (r2 range for the best fit models = 0.27-0.36). At neighborhood scales, species spatial associations were positively correlated with similarity in π(tlp), consistent with predictions for hierarchical competition. Although the correlation between π(tlp) and interspecific spatial associations was weak (r2 < 0.01), this showed a persistent influence of drought tolerance on neighborhood interactions and community assembly. Quantifying the full impact of traits on competitive interactions in forests may require incorporating plasticity among individuals within species, especially among specific life stages, and moving beyond individual traits to integrate the impact of multiple traits on whole-plant performance and resource demand.

This data paper provides a description of OpenNahele, the open Hawaiian forest plot database. OpenNahele includes 530 forest plots across the Hawaiian archipelago containing 43,590 individuals of 185 native and alien tree, shrub and tree fern species across six islands. We include estimates of maximum plant size (D950.1 and Dmax3) for 58 woody plant species, a key functional trait associated with dispersal distance and competition for light. OpenNahele can serve as a platform to test key ecological, evolutionary and conservation questions in a hotspot archipelago. OpenNahele is the first database that compiles data from a large number of forest plots across the Hawaiian archipelago to allow broad and high resolution studies of biodiversity patterns. Keywords: Hawaii, forests, islands, biodiversity, community ecology, evolutionary ecology

Environmental filtering and dispersal limitation can both maintain diversity in plant communities by aggregating conspecifics, but parsing the contribution of each process has proven difficult empirically. Here, we assess the contribution of filtering and dispersal limitation to the spatial aggregation patterns of 456 tree species in a hyperdiverse Amazonian forest and find distinct functional trait correlates of interspecific variation in these processes. Spatial point process model analysis revealed that both mechanisms are important drivers of intraspecific aggregation for the majority of species. Leaf drought tolerance was correlated with species topographic distributions in this aseasonal rainforest, showing that future increases in drought severity could significantly impact community structure. In addition, seed mass was associated with the spatial scale and density of dispersal-related aggregation. Taken together, these results suggest environmental filtering and dispersal limitation act in concert to influence the spatial and functional structure of diverse forest communities.

As the range of studies on macroecology and functional traits expands, integration of traits into higher-level approaches offers new opportunities to improve clarification of larger-scale patterns and their mechanisms and predictions using models. Here, we propose a framework for quantifying ‘ecosystem traits’ and means to address the challenges of broadening the applicability of functional traits to macroecology. Ecosystem traits are traits or quantitative characteristics of organisms (plants, animals, and microbes) at the community level expressed as the intensity (or density) normalized per unit land area. Ecosystem traits can inter-relate and integrate data from field trait surveys, eddy-flux observation, remote sensing, and ecological models, and thereby provide new resolution of the responses and feedback at regional to global scale.

Aim: To determine the role of regional forcing on plot-level species diversity and composition, and to quantify the relative importance of biogeographical and climatic factors in explaining woody plant diversity and composition at the local-, island- and archipelago-scale. Location: Forty-one tropical islands of the Indo-Pacific region from Madagascar to Hawai‘i Island. Methods: We analysed the diversity and composition of tropical woody plant communities located across 113 plots, 41 islands and 19 archipelagos. We used generalized linear mixed-effects models and generalized dissimilarity models to determine the role of regional forcing at the island and archipelago scale and to assess the relative importance of biogeographical (area and isolation of islands or archipelagos, geographical distance between plots) and climatic factors in explaining differences in local diversity and composition (species turnover). Analyses were conducted at different geographical scales (local, island and archipelago) and taxonomic levels (species, genus and family). Results: Variation in local (plot-level) diversity (as species density, the number of species per 100 woody plants) was primarily explained by island and archipelago identity. Maximum species density was positively correlated with the area of an island (or archipelago) and negatively correlated with the isolation of an archipelago. Local climatic variability was also a significant predictor of species density, but less important than regional forcing. Climate variables explained < 20% of the variation in species turnover across all plots. The importance of geographical distance between plots relative to climate in driving species turnover decreased from the species to family level, and from the regional to island level. Main conclusions: Regional forcing was the key driver of local diversity and composition on islands. Island area and archipelago isolation are likely driving local diversity through their effects on the pool of island species. Geographical distance between plots is the main factor explaining species turnover, while at higher taxonomic levels, climatic factors and niche conservatism are the main drivers. Refereed/Peer-reviewed

The need for rigorous analyses of climate impacts has never been more crucial. Current textbooks state that climate directly influences ecosystem annual net primary productivity (NPP), emphasizing the urgent need to monitor the impacts of climate change. A recent paper challenged this consensus, arguing, based on an analysis of NPP for 1247 woody plant communities across global climate gradients, that temperature and precipitation have negligible direct effects on NPP and only perhaps have indirect effects by constraining total stand biomass (Mtot ) and stand age (a). The authors of that study concluded that the length of the growing season (lgs ) might have a minor influence on NPP, an effect they considered not to be directly related to climate. In this article, we describe flaws that affected that study's conclusions and present novel analyses to disentangle the effects of stand variables and climate in determining NPP. We re-analyzed the same database to partition the direct and indirect effects of climate on NPP, using three approaches: maximum-likelihood model selection, independent-effects analysis, and structural equation modeling. These new analyses showed that about half of the global variation in NPP could be explained by Mtot combined with climate variables and supported strong and direct influences of climate independently of Mtot , both for NPP and for net biomass change averaged across the known lifetime of the stands (ABC = average biomass change). We show that lgs is an important climate variable, intrinsically correlated with, and contributing to mean annual temperature and precipitation (Tann and Pann ), all important climatic drivers of NPP. Our analyses provide guidance for statistical and mechanistic analyses of climate drivers of ecosystem processes for predictive modeling and provide novel evidence supporting the strong, direct role of climate in determining vegetation productivity at the global scale.

Leaf hydraulic conductance (Kleaf ) quantifies the capacity of a leaf to transport liquid water and is a major constraint on light-saturated stomatal conductance (gs ) and photosynthetic rate (Amax ). Few studies have tested the plasticity of Kleaf and anatomy across growth light environments. These provided conflicting results. The Hawaiian lobeliads are an excellent system to examine plasticity, given the striking diversity in the light regimes they occupy, and their correspondingly wide range of Amax , allowing maximal carbon gain for success in given environments. We measured Kleaf , Amax , gs and leaf anatomical and structural traits, focusing on six species of lobeliads grown in a common garden under two irradiances (300/800 μmol photons m(-2) s(-1) ). We tested hypotheses for light-induced plasticity in each trait based on expectations from optimality. Kleaf , Amax , and gs differed strongly among species. Sun/shade plasticity was observed in Kleaf , Amax, and numerous traits relating to lamina and xylem anatomy, venation, and composition, but gs was not plastic with growth irradiance. Species native to higher irradiance showed greater hydraulic plasticity. Our results demonstrate that a wide set of leaf hydraulic, stomatal, photosynthetic, anatomical, and structural traits tend to shift together during plasticity and adaptation to diverse light regimes, optimizing performance from low to high irradiance.

Resolving the drivers of hydraulic decline during drought is crucial for understanding drought tolerance in crops and natural ecosystems. In the past 15 years, studies of the decline of leaf hydraulic conductance (Kleaf) have supported a major role in controlling plant drought responses. We analyzed the variation in Kleaf decline with dehydration in a global database of 310 species, providing novel insights into its underlying mechanisms, its co-ordination with stem hydraulics, its influence on gas exchange and drought tolerance, and its linkage with species ecological distributions. Kleaf vulnerability varied strongly within and across lineages, growth forms, and biomes. A critical literature review indicates that changes in hydraulic conductance outside the xylem with dehydration drive the overall decline of Kleaf. We demonstrate a significant leaf hydraulic safety-efficiency trade-off across angiosperm species and discuss the importance of the large variation around this trend. Leaves tend to be more vulnerable than stems, with their vulnerabilities co-ordinated across species, and importantly linked with adaptation across biomes. We hypothesize a novel framework to explain diversity across species in the co-ordination of Kleaf and gas exchange during dehydration. These findings reflect considerable recent progress, yet new tools for measurement, visualization, and modeling will result in ongoing discoveries important across fields in plant biology.

Altres ajuts: this research was partially funded by a Natural Sciences and Engineering Research Grant to BS. VO was supported by the Russian Science Foundation (#14-50-00029). The study has been supported by theTRY initiative on plant traits (http://www.try-db.org).The TRY initiative and database is hosted at the MaxPlanck Institute for Biogeochemistry, Jena, Germany.TRY is currently supported by DIVERSITAS/FutureEarth and the German Centre for Integrative Biodiver-sity Research (iDiv) Halle-Jena-Leipzig. Questions: Heinz Ellenberg classically defined "indicator" scores for species representing their typical positions along gradients of key environmental variables, and these have proven very useful for designating ecological distributions. We tested a key tenent of trait-based ecology, i.e. the ability to predict ecological preferences from species' traits. More specifically, can we predict Ellenberg indicator scores for soil nutrients, soil moisture and irradiance from four well-studied traits: leaf area, leaf dry matter content, specific leaf area (SLA) and seed mass? Can we use such relationships to estimate Ellenberg scores for species never classified by Ellenberg? - Location: global. - Methods: cumulative link models were developed to predict Ellenberg nutrients, irradiance and moisture values from Ln-transformed trait values using 922, 981 and 988 species, respectively. We then independently tested these prediction equations using the trait values of 423 and 421 new species that occurred elsewere in Europe, North America and Morocco, and whose habitat affinities we could classify from independent sources as three-level ordinal ranks related to soil moisture and irradiance. The traits were SLA, leaf dry matter content, leaf area and seed mass. - Results: the four functional traits predicted the Ellenberg indicator scores of site fertility, light and moisture with average error rates of

Pan-tropically, liana density increases with decreasing rainfall and increasing seasonality. This pattern has led to the hypothesis that lianas display a growth advantage over trees under dry conditions. However, the physiological mechanisms underpinning this hypothesis remain elusive. A key trait influencing leaf and plant drought tolerance is the leaf water potential at turgor loss point (πtlp).πtlpadjusts under drier conditions and this contributes to improved leaf drought tolerance. For co-occurring Amazonian tree (n= 247) and liana (n= 57) individuals measured during the dry and the wet seasons, lianas showed a stronger osmotic adjustment than trees. Liana leaves were less drought-tolerant than trees in the wet season, but reached similar drought tolerances during the dry season. Stronger osmotic adjustment in lianas would contribute to turgor maintenance, a critical prerequisite for carbon uptake and growth, and to the success of lianas relative to trees in growth under drier conditions.

Many species face increasing drought under climate change. Plasticity has been predicted to strongly influence species' drought responses, but broad patterns in plasticity have not been examined for key drought tolerance traits, including turgor loss or 'wilting' point (πtlp ). As soil dries, plants shift πtlp by accumulating solutes (i.e. 'osmotic adjustment'). We conducted the first global analysis of plasticity in Δπtlp and related traits for 283 wild and crop species in ecosystems worldwide. Δπtlp was widely prevalent but moderate (-0.44 MPa), accounting for 16% of post-drought πtlp. Thus, pre-drought πtlp was a considerably stronger predictor of post-drought πtlp across species of wild plants. For cultivars of certain crops Δπtlp accounted for major differences in post-drought πtlp. Climate was correlated with pre- and post-drought πtlp, but not Δπtlp. Thus, despite the wide prevalence of plasticity, πtlp measured in one season can reliably characterise most species' constitutive drought tolerances and distributions relative to water supply.

Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25ha), all stems 1cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 degrees S-61 degrees N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 degrees C), changes in precipitation (up to +/- 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8g Nm(-2)yr(-1) and 3.1g Sm(-2)yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.

For niche differences to maintain coexistence of sympatric species, each species must grow and/or survive better than each of the others in at least one set of conditions (i.e., performance trade-offs). However, the extent of niche differentiation in tropical forests remains highly debated. We present the first test of performance trade-offs for wild seedlings in a tropical forest. We measured seedling relative growth rate (RGR) and survival of four common native woody species across 18 light, substrate, and topography microhabitats over 2.5 years within Hawaiian montane wet forest, an ideal location due to its low species diversity and strong species habitat associations. All six species pairs exhibited significant performance trade-offs across microhabitats and for RGR versus survival within microhabitats. We also found some evidence of performance equivalence, with species pairs having similar performance in 26% of comparisons across microhabitats. Across species, survival under low light was generally positively associated with RGR under high light. When averaged over all species, topography (slope, aspect, and elevation) explained most of the variation in RGR attributable to microhabitat variables (51-53%) followed by substrate type (35-37%) and light (11-12%). However, the relative effects of microhabitat differed among species and RGR metric (i.e., RGR for height, biomass, or leaf area). These findings indicate that performance trade-offs among species during regeneration are common in low-diversity tropical forest, although other mechanisms may better explain the coexistence of species with small performance differences.

Summary 1. The leaf economics spectrum (LES) has been an organizing framework of plant functional ecology for the past decade. The LES describes a set of trade-offs among traits related to plant carbon balance. Species with a long leaf life span (LLS) invest additional material for leaf protection and structural support and consequently tend to have a lower leaf photosynthetic rate per unit mass than species with a shorter LLS. 2. While the LES is most apparent in comparing species with extreme differences in their traits, it has nonetheless been adopted as a general explanation of leaf trait variation at all scales and in all plants. It highlights the ‘trait-based’ approach to plant ecology, which has generally used a small set of traits to predict whole organism and even whole ecosystem attributes. Few studies have investigated the relationships between LES traits and organismal attributes not directly related to carbon economy. 3. We explored the LES in 32 deciduous woody species of Viburnum (Adoxaceae). We found no evidence for any mass-based LES trade-offs. Rather, on an area basis, photosynthetic rates were positively correlated with leaf mass per area (LMA); higher LMA was associated with greater investment in photosynthetic tissue, with most of the variation due to changes in the thickness of photosynthetic mesophyll. 4. Species’ mean LLS varied between 19 and 26 weeks and was not correlated with other LES traits. Instead, LLS was strongly associated with the diverse set of whole-plant branching patterns in Viburnum. In the most common growth pattern, LLS was significantly correlated with flowering time, because branches end in terminal inflorescences, and all leaves and inflorescences are preformed in overwintering buds. 5. Synthesis. Plants may recover the cost of their leaves early in the growing season, allowing LLS to vary independently of the plant carbon budget. In deciduous species, LLS may be strongly influenced by whole plant architecture, which, in Viburnum, is evolutionarily conserved. In general, positive area-based LES trait relationships will limit the relevance of LLS to this spectrum and allow LLS to vary for reasons that are not directly related to carbon economy.

Question: Are plant traits more closely correlated with mean annual temperature, orwithmean annual precipitation? Location: Global. Methods: We quantified the strength of the relationships between temperature and precipitation and 21 plant traits from 447,961 species-site combinations worldwide. We used meta-analysis to provide an overall answer to our question. Results: Mean annual temperature was significantly more strongly correlated with plant traits than was mean annual precipitation. Conclusions: Our study provides support for some of the assumptions of classical vegetation theory, and points to many interesting directions for future research. The relatively low R

Summary The mechanical resistance of leaves has key ecological implications but its basis has not been well understood, particularly at the tissue scale. We tested the hypotheses that leaf mechanical resistance should be a function of tissue density, increasing from the lamina to the midrib, and higher in drought-tolerant than drought-avoiding species. In a common garden study, we quantified nine leaf biomechanical traits, including measurements of material and structural resistance, and in addition 17 morphological traits, at the tissue and whole-leaf scales, for 21 species from three semi-arid communities of California, USA. The mechanical properties of leaves depended strongly on tissue density. Material resistance was significantly greater in the midrib than in the leaf lamina, and tissue resistances were significantly correlated among tissues, lower in deciduous coastal sage species and higher in evergreen drought-tolerant chaparral species. The proportion of the biomass invested in the midrib was lower in species bearing midribs and laminas of high material resistance. Our results support the hypothesis of a hierarchical partitioning of leaf mechanical resistance among leaf tissues reflecting the investment of dry mass. Also, our data indicated a mechanical compensation in leaf design, where leaves with high material resistance and density deploy a relatively minor proportion of support tissue in the midrib. Finally, our results establish a quantitative basis for differences among communities in leaf biomechanics. Our results supported the classical characterization of the mediterranean-climate flora of California according to the dramatic increase in the mean leaf mechanical resistance from species of coastal sage to chaparral, with diverse leaf types in the Mojave Desert species.

Pérez-Harguindeguy et al., Plant functional traits are the features (morphological, physiological, phenological) that represent ecological strategies and determine how plants respond to environmental factors, affect other trophic levels and influence ecosystem properties. Variation in plant functional traits, and trait syndromes, has proven useful for tackling many important ecological questions at a range of scales, giving rise to a demand for standardised ways to measure ecologically meaningful plant traits. This line of research has been among the most fruitful avenues for understanding ecological and evolutionary patterns and processes. It also has the potential both to build a predictive set of local, regional and global relationships between plants and environment and to quantify a wide range of natural and human-driven processes, including changes in biodiversity, the impacts of species invasions, alterations in biogeochemical processes and vegetation-atmosphere interactions. The importance of these topics dictates the urgent need for more and better data, and increases the value of standardised protocols for quantifying trait variation of different species, in particular for traits with power to predict plant-and ecosystem-level processes, and for traits that can be measured relatively easily. Updated and expanded from the widely used previous version, this handbook retains the focus on clearly presented, widely applicable, step-by-step recipes, with a minimum of text on theory, and not only includes updated methods for the traits previously covered, but also introduces many new protocols for further traits. This new handbook has a better balance between whole-plant traits, leaf traits, root and stem traits and regenerative traits, and puts particular emphasis on traits important for predicting species' effects on key ecosystem properties. We hope this new handbook becomes a standard companion in local and global efforts to learn about the responses and impacts of different plant species with respect to environmental changes in the present, past and future. © CSIRO 2013., This contribution was funded, in part, by the Inter-American Institute for Global Change Research (IAI) CRN 2015 and SGP-CRA2015, which were supported by US National Science Foundation Grants GEO-0452325 and GEO-1138881 The work was also funded by FONCyT (PICT 20441 and 365), CONICET (PIP 2006-2011 11220080101532), SECYT-Universidad Nacional de Córdoba – Argentina, and the US NSF (DEB-0620652).

© 2017 John Wiley & Sons Ltd/CNRS Leaf dry mass per unit leaf area (LMA) is a central trait in ecology, but its anatomical and compositional basis has been unclear. An explicit mathematical and physical framework for quantifying the cell and tissue determinants of LMA will enable tests of their influence on species, communities and ecosystems. We present an approach to explaining LMA from the numbers, dimensions and mass densities of leaf cells and tissues, which provided unprecedented explanatory power for 11 broadleaved woody angiosperm species diverse in LMA (33–262 g m−2; R2 = 0.94; P

AimThe taxon cycle hypothesis describes the cyclic movement of taxa during range expansion and contraction, accompanied by an evolutionary shift from open and often coastal vegetation to closed, and often inland forest vegetation in island systems. The Hawaiian Archipelago is an ideal system to test this hypothesis given the linear fashion of island formation and a relatively well-understood geological history.LocationHawaiian Islands.MethodsWe sampled 153 individuals in 15 of the 16 native species of HawaiianEuphorbiasectionAnisophyllumon six major Hawaiian Islands, plus 11 New World close relatives, to elucidate the biogeographic movement of this lineage along the Hawaiian island chain. We used a concatenated chloroplast DNA data set of more than eight kilobases in aligned length and applied maximum likelihood and Bayesian inference for phylogenetic reconstruction. Connectivity among islands and habitat types was estimated using BayesTraits. Age and phylogeographic patterns were co-estimated using BEAST. In addition, we used nuclear ribosomal ITS and the low-copy genesLEAFYandG3pdhCto investigate the reticulate relationships within this radiation.ResultsWe estimate that HawaiianEuphorbiafirst arrived on Kauai or Niihau ca. 5 million years ago and subsequently diverged into 16 species on all major Hawaiian Islands. During this processEuphorbiadispersed from older to younger islands in a stepping-stone fashion through open, dispersal-prone habitats. Taxa that occupy closed vegetation on Kauai and Oahu evolvedin situfrom open vegetation taxa of the same island. Consequently, widespread species tend to occupy habitats with open vegetation, whereas single island endemic species predominantly occur in habitats with closed canopy and are only found on the two oldest islands of Kauai and Oahu.Main conclusionsThe spatial and temporal patterns of dispersal and range shifts in HawaiianEuphorbiasupport an intra-volcanic-archipelago version of the taxon cycle hypothesis.

© 2016 John Wiley & Sons Ltd It was shown over 40 years ago that plants maximize carbon gain for a given rate of water loss if stomatal conductance, gs, varies in response to external and internal conditions such that the marginal carbon revenue of water, ∂A/∂E, remains constant over time. This theory has long held promise for understanding the physiological ecology of water use and for informing models of plant-atmosphere interactions. Full realization of this potential hinges on three questions: (i) Are analytical approximations adequate for applying the theory at diurnal time scales? (ii) At what time scale is it realistic and appropriate to apply the theory? (iii) How should gs vary to maximize growth over long time scales? We review the current state of understanding for each of these questions and describe future research frontiers. In particular, we show that analytical solutions represent the theory quite poorly, especially when boundary layer or mesophyll resistances are significant; that diurnal variations in hydraulic conductance may help or hinder maintenance of ∂A/∂E, and the matter requires further study; and that optimal diurnal responses are distinct from optimal long-term variations in gs, which emerge from optimal shifts in carbon partitioning at the whole-plant scale.

International audience; Amazonian tree communities have already been seriously impacted by extreme natural droughts, and intense droughts are predicted to increase in frequency. However, our current knowledge of Amazonian tree species' responses to water stress remains limited, as plant trait databases include few drought tolerance traits, impeding the application and predictive power of models. Here we explored how leaf water potential at turgor loss point (π tlp), a determinant of leaf drought tolerance, varies with species life history, season, tree size and irradiance within a forest in French Guiana. First, we provided a further direct validation of a rapid method of π tlp determination based on osmometer measurements of leaf osmotic potential at full hydration for five Amazonian tree species. Next, we analysed a dataset of 131 π tlp values for a range of species, seasons, size (including saplings), and leaf exposure. We found that early-successional species had less drought-tolerant leaves than late-successional species. Species identity was the major driver of π tlp variation, whereas season, canopy tree size and leaf exposure explained little variation. Shifts in π tlp from saplings to canopy trees varied across species, and sapling leaf drought tolerance was a moderate predictor of canopy tree leaf drought tolerance. Given its low within-species variability, we propose that π tlp is a robust trait, and is useful as one index of species' drought tolerance. We also suggest that measuring this trait would considerably advance our knowledge on leaf drought tolerance in hyperdiverse communities and would thus likely shed light on the resilience of such vulnerable species-rich ecosystem.

Water plays a central role in plant biology and the efficiency of water transport throughout the plant affects both photosynthetic rate and growth, an influence that scales up deterministically to the productivity of terrestrial ecosystems. Moreover, hydraulic traits mediate the ways in which plants interact with their abiotic and biotic environment. At landscape to global scale, plant hydraulic traits are important in describing the function of ecological communities and ecosystems. Plant hydraulics is increasingly recognized as a central hub within a network by which plant biology is connected to palaeobiology, agronomy, climatology, forestry, community and ecosystem ecology and earth-system science. Such grand challenges as anticipating and mitigating the impacts of climate change, and improving the security and sustainability of our food supply rely on our fundamental knowledge of how water behaves in the cells, tissues, organs, bodies and diverse communities of plants. A workshop, 'Emerging Frontiers in Plant Hydraulics' supported by the National Science Foundation, was held in Washington DC, 2015 to promote open discussion of new ideas, controversies regarding measurements and analyses, and especially, the potential for expansion of up-scaled and down-scaled inter-disciplinary research, and the strengthening of connections between plant hydraulic research, allied fields and global modelling efforts.

Strong latitudinal patterns in leaf form are well documented in floristic comparisons and palaeobotanical studies. However, there is little agreement about their functional significance; in fact, it is still unknown to what degree these patterns were generated by repeated evolutionary adaptation. We analysed leaf form in the woody angiosperm clade Viburnum (Adoxaceae) and document evolutionarily correlated shifts in leafing habit, leaf margin morphology, leaf shape and climate. Multiple independent shifts between tropical and temperate forest habitats have repeatedly been accompanied by a change between evergreen, elliptical leaves with entire margins and deciduous, more rounded leaves with toothed or lobed margins. These consistent shifts in Viburnum support repeated evolutionary adaptation as a major determinant of the global correlation between leaf form and mean annual temperature. Our results provide a new theoretical grounding for the inference of past climates using fossil leaf assemblages.