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Variation in the architecture of trait networks among ecosystems has been rarely quantified, but can provide high resolution of the contrasting adaptation of the whole phenotype. We constructed leaf trait networks (LTNs) from 35 structural, anatomical and compositional leaf traits for 394 tree species in nine forests from tropical to cold-temperate zones in China. Our analyses supported the hypothesis that LTNs would increase in modular complexity across forests in parallel with species-richness and climatic warmth and moisture, due to reduced phenotypic constraints and greater opportunities for niche differentiation. Additionally, we found that within LTNs, leaf economics traits including leaf thickness would have central importance, acting as hub traits with high connectivity due to their contributions to multiple functions. Across the continent, the greater species richness and trait diversity observed in forests under resource-rich climates enable greater complexity in whole phenotype structure and function as indicated by the trait network architecture.

Considering the global intensification of aridity in tropical biomes due to climate change, we need to understand what shapes the distribution of drought sensitivity in tropical plants. We conducted a pantropical data synthesis representing 1117 species to test whether xylem-specific hydraulic conductivity (KS), water potential at leaf turgor loss (ΨTLP) and water potential at 50% loss of KS (ΨP50) varied along climate gradients. The ΨTLP and ΨP50 increased with climatic moisture only for evergreen species, but KS did not. Species with high ΨTLP and ΨP50 values were associated with both dry and wet environments. However, drought-deciduous species showed high ΨTLP and ΨP50 values regardless of water availability, whereas evergreen species only in wet environments. All three traits showed a weak phylogenetic signal and a short half-life. These results suggest strong environmental controls on trait variance, which in turn is modulated by leaf habit along climatic moisture gradients in the tropics., Ecology Letters, 25 (12), ISSN:1461-023X, ISSN:1461-0248

PREMISE Fossil leaf traits can enable reconstruction of ancient environments and climates. Among these, leaf size has been particularly studied because it reflects several climatic forcings (e.g., precipitation and surface temperature) and, potentially, environment characteristics (e.g., nutrient availability, local topography, and openness of vegetation). However, imperfect preservation and fragmentation can corrupt its utilization. We provide improved methodology to estimate leaf size from fossil fragments. METHODS We apply three methods: (1) visually reconstructing leaf area based on taxon-specific gross morphology; (2) estimating intact leaf area from vein density based on a vein scaling relationship; and (3) a novel complementary method, determining intact leaf length based on the tapering of the midvein in the fragment. We test the three methods for fossils of extinct Eotrigonobalanus furcinervis (Fagaceae) from two lignite horizons of the middle and late Eocene of central Germany respectively (~45/46 and 35/36 Ma). RESULTS The three methods, including the new one, yield consistent leaf size reconstructions. The vein scaling method showed a shift to larger leaf size, from the middle to the late Eocene. CONCLUSIONS These methods constitute a toolbox with different solutions to reconstruct leaf size from fossil fragments depending on fossil preservation. Fossil leaf size reconstruction has great potential to improve physiognomy-based paleoenvironmental reconstructions and the interpretation of the fossil record.

Functional traits are frequently used to evaluate plant adaptation across environments. Yet, traits tend to have multiple functions and interactions, which cannot be accounted for in traditional correlation analyses. Plant trait networks (PTNs) clarify complex relationships among traits, enable the calculation of metrics for the topology of trait coordination and the importance of given traits in PTNs, and how they shift across communities. Recent studies of PTNs provide new insights into some important topics, including trait dimensionality, trait spectra (including the leaf economic spectrum), stoichiometric principles, and the variation of phenotypic integration along gradients of resource availability. PTNs provide improved resolution of the multiple dimensions of plant adaptation across scales and responses to shifting resources, disturbance regimes, and global 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.

International audience; The links between internal nitrogen recycling through the process of resorption from senescing leaves, whole-plant resource-use strategies and performance remain elusive. Indeed, tests of such potential linkages are hampered by the classical evaluation of plant nitrogen resorption efficiency (REN) based on ‘snapshots’ of leaf nitrogen concentration from adult and senescent leaves. We significantly increased the resolution for measuring leaf nitrogen resorption by non-destructively tracking time courses of nitrogen concentration in leaves of 137 natural Arabidopsis thaliana genotypes native to a wide range of climates across Europe, grown in a greenhouse. In addition to the classical measurement of resorption efficiency, we computed the relative maximum resorption rate of nitrogen (RRN), that is, the amount of nitrogen remobilized by a leaf per unit time and per unit nitrogen, together with slow–fast syndrome traits at the leaf and whole-plant levels. Across genotypes, high rates and efficiencies of nitrogen resorption were associated with low specific leaf area and late flowering. The RRN showed significant heritability, genetic associations and was negatively correlated with the mean annual temperature of the native populations. By contrast, despite the evidence of a correlation with temperature, REN showed lower heritability and no evidence of genetic association, questioning the mechanisms of nitrogen resorption. Overall, our results suggest a much stronger adaptive role for leaf nitrogen resorption than previously uncovered.

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.

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.

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.

The application of functional traits to predict and explain plant species’ distributions and vital rates has been a major direction in functional ecology for decades, yet numerous physiological traits have not yet been incorporated into the approach. Using commonly measured traits such as leaf mass per area (LMA) and wood density (WD), and additional traits related to water transport, gas exchange and resource economics, including leaf vein, stomatal and wilting traits, we tested hypotheses for Hawaiian wet montane and lowland dry forests (MWF and LDF, respectively): (1) Forests would differ in a wide range of traits as expected from contrasting adaptation; (2) trait values would be more convergent among dry than wet forest species due to the stronger environmental filtering; (3) traits would be intercorrelated within “modules” supporting given functions; (4) relative growth rate (RGR) and mortality rate (m) would correlate with a number of specific traits; with (5) stronger relationships when stratifying by tree size; and (6) RGR and m can be strongly explained from trait‐based models. The MWF species’ traits were associated with adaptation to high soil moisture and nutrient supply and greater shade tolerance, whereas the LDF species’ traits were associated with drought tolerance. Thus, on average, MWF species achieved higher maximum heights than LDF species and had leaves with larger epidermal cells, higher maximum stomatal conductance and CO2 assimilation rate, lower vein lengths per area, higher saturated water content and greater shrinkage when dry, lower dry matter content, higher phosphorus concentration, lower nitrogen to phosphorus ratio, high chlorophyll to nitrogen ratio, high carbon isotope discrimination, high stomatal conductance to nitrogen ratio, less negative turgor loss point and lower WD. Functional traits were more variable in the MWF than LDF, were correlated within modules, and predicted species’ RGR and m across forests, with stronger relationships when stratifying by tree size. Models based on multiple traits predicted vital rates across forests (R2 = 0.70–0.72; p

ForestGEO is a network of scientists and long-term forest dynamics plots (FDPs) spanning the Earth's major forest types. ForestGEO's mission is to advance understanding of the diversity and dynamics of forests and to strengthen global capacity for forest science research. ForestGEO is unique among forest plot networks in its large-scale plot dimensions, censusing of all stems ≥1 cm in diameter, inclusion of tropical, temperate and boreal forests, and investigation of additional biotic (e.g., arthropods) and abiotic (e.g., soils) drivers, which together provide a holistic view of forest functioning. The 71 FDPs in 27 countries include approximately 7.33 million living trees and about 12,000 species, representing 20% of the world's known tree diversity. With >1300 published papers, ForestGEO researchers have made significant contributions in two fundamental areas: species coexistence and diversity, and ecosystem functioning. Specifically, defining the major biotic and abiotic controls on the distribution and coexistence of species and functional types and on variation in species' demography has led to improved understanding of how the multiple dimensions of forest diversity are structured across space and time and how this diversity relates to the processes controlling the role of forests in the Earth system. Nevertheless, knowledge gaps remain that impede our ability to predict how forest diversity and function will respond to climate change and other stressors. Meeting these global research challenges requires major advances in standardizing taxonomy of tropical species, resolving the main drivers of forest dynamics, and integrating plot-based ground and remote sensing observations to scale up estimates of forest diversity and function, coupled with improved predictive models. However, they cannot be met without greater financial commitment to sustain the long-term research of ForestGEO and other forest plot networks, greatly expanded scientific capacity across the world's forested nations, and increased collaboration and integration among research networks and disciplines addressing forest science.

[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.

© 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.

Woody perennial plants on islands have repeatedly evolved from herbaceous mainland ancestors. Although the majority of species in Euphorbia subgenus Chamaesyce section Anisophyllum (Euphorbiaceae) are small and herbaceous, a clade of 16 woody species diversified on the Hawaiian Islands. They are found in a broad range of habitats, including the only known C4 plants adapted to wet forest understories. We investigate the history of island colonization and habitat shift in this group. We sampled 153 individuals in 15 of the 16 native species of Hawaiian Euphorbia on six major Hawaiian Islands, plus 11 New World close relatives, to elucidate the biogeographic movement of this lineage within 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. Age and phylogeographic patterns were co‐estimated using BEAST. In addition, we used nuclear ribosomal ITS and the low‐copy genes LEAFY and G3pdhC to investigate the reticulate relationships within this radiation. Hawaiian Euphorbia first arrived on Kaua`i or Ni`ihau ca. 5 million years ago and subsequently diverged into 16 named species with extensive reticulation. During this process Hawaiian Euphorbia dispersed from older to younger islands through open vegetation that is disturbance‐prone. Species that occur under closed vegetation evolved in situ from open vegetation of the same island and are only found on the two oldest islands of Kaua`i and O`ahu. The biogeographic history of Hawaiian Euphorbia supports a progression rule with within‐island shifts from open to closed vegetation.

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.

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.

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.

PREMISE OF THE STUDY Studies across diverse species have established theory for the contribution of leaf traits to plant drought tolerance. For example, species in more arid climates tend to have smaller leaves of higher vein density, higher leaf mass per area, and more negative osmotic potential at turgor loss point (πTLP ). However, few studies have tested these associations for species within a given lineage that have diversified across an aridity gradient. METHODS We analyzed the anatomy and physiology of 10 Ceanothus (Rhamnaceae) species grown in a common garden for variation between and within "wet" and "dry" subgenera (Ceanothus and Cerastes, respectively) and analyzed a database for 35 species for leaf size and leaf mass per area (LMA). We used a phylogenetic generalized least squares approach to test hypothesized relationships among traits, and of traits with climatic aridity in the native range. We also tested for allometric relationships among anatomical traits. KEY RESULTS Leaf form, anatomy, and drought tolerance varied strongly among species within and between subgenera. Cerastes species had specialized anatomy including hypodermis and encrypted stomata that may confer superior water storage and retention. The osmotic potentials at turgor loss point (πTLP ) and full turgor (πo ) showed evolutionary correlations with the aridity index (AI) and precipitation of the 10 species' native distributions, and LMA with potential evapotranspiration for the 35 species in the larger database. We found an allometric correlation between upper and lower epidermal cell wall thicknesses, but other anatomical traits diversified independently. CONCLUSIONS Leaf traits and drought tolerance evolved within and across lineages of Ceanothus consistently with climatic distributions. The πTLP has signal to indicate the evolution of drought tolerance within small clades.

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

1. Water availability is a key determinant of forest ecosystem function and tree species distributions. While droughts are increasing in frequency in many ecosystems, including in the tropics, plant responses to water supply vary with species and drought intensity and are therefore difficult to model. Based on physiological first principles, we hypothesized that trees with a lower turgor loss point (pi(tlp)), that is, a more negative leaf water potential at wilting, would maintain water transport for longer into a dry season. 2. We measured sapflux density of 22 mature trees of 10 species during a dry season in an Amazonian rainforest, quantified sapflux decline as soil water content decreased and tested its relationship to tree pi(tlp), size and leaf predawn and midday water potentials measured after the onset of the dry season. 3. The measured trees varied strongly in the response of water use to the seasonal drought, with sapflux at the end of the dry season ranging from 37 to 117% (on average 83 +/- 5 %) of that at the beginning of the dry season. The decline of water transport as soil dried was correlated with tree pi(tlp) (Spearman's rho >= 0.63), but not with tree size or predawn and midday water potentials. Thus, trees with more drought-tolerant leaves better maintained water transport during the seasonal drought. 4. Our study provides an explicit correlation between a trait, measurable at the leaf level, and whole-plant performance under drying conditions. Physiological traits such as pi(tlp) can be used to assess and model higher scale processes in response to drying 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.

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.

Premise of research. Leaf mesophyll is often differentiated into a palisade layer with tightly packed, elongated cells (I-cells) and a spongy layer with loosely packed, complex shaped cells. An alternative palisade type, composed of branched H-cells, has evolved in a number of plant lineages. Viburnum (Adoxaceae) possesses both types of palisade, providing an opportunity to assess the significance of evolutionary switches between these forms.Methodology. An anatomical survey of 80 species spanning the Viburnum phylogeny permitted an analysis of palisade differences in relation to other characters. A geometric model of leaf mesophyll surface area for CO2 absorption correlated well with measured photosynthetic capacity in a subset of species, allowing us to infer shifts in photosynthetic function.Pivotal results. Ancestrally, viburnums probably produced a palisade with one layer of H-cells. Multiple transitions to two layers of H-cells (H2) and to one or two layers of I-cells (I1, I2) occurred. These shifts...

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

1. Across plant species, drought tolerance and distributions with respect to water availability are strongly correlated with two physiological traits, the leaf water potential at wilting, that is, turgor loss point (ptlp), and the cell solute potential at full hydration, that is, osmotic potential (po). We present methods to determine these parameters 30 times more rapidly than the standard pressurevolume (pv) curve approach, making feasible community-scale studies of plant drought tolerance. 2. We optimized existing methods for measurements of pi o using vapour-pressure osmometry of freeze-thawed leaf discs from 30 species growing in two precipitation regimes, and developed the first regression relationships to accurately estimate pressurevolume curve values of both pi o and pi tlp from osmometer values. 3. The pi o determined with the osmometer (pi osm) was an excellent predictor of the pi o determined from the pv curve (pi pv,r2 = 0.80). Although the correlation of pi osm and pi pv enabled prediction, the relationship departed from the 1 : 1 line. The discrepancy between the methods could be quantitatively accounted for by known sources of error in osmometer measurements, that is, dilution by the apoplastic water, and solute dissolution from destroyed cell walls. An even stronger prediction of pi pv could be made using pi osm, leaf density (rho), and their interaction (r2 = 0.85, all P l 2 x 10-10). 4. The pi osm could also be used to predict pi tlp (r2 = 0.86). Indeed, pi osm was a better predictor of pi tlp than leaf mass per unit area (LMA; r2 = 0.54), leaf thickness (T; r2 = 0.12), rho (r2 = 0.63), and leaf dry matter content (LDMC; r2 = 0.60), which have been previously proposed as drought tolerance indicators. Models combining posm with LMA, T, rho, or LDMC or other pv curve parameters (i.e. elasticity and apoplastic fraction) did not significantly improve prediction of pi tlp. 5. This osmometer method enables accurate measurements of drought tolerance traits across a wide range of leaf types and for plants with diverse habitat preferences, with a fraction of effort of previous methods. We expect it to have wide application for predicting species responses to climate variability and for assessing ecological and evolutionary variation in drought tolerance in natural populations and agricultural cultivars.

1. Xylem hydraulic properties play an essential role in supporting growth and photosynthesis and influence sensitivity to environmental conditions such as drought and freezing. Consequently, stem hydraulic conductance can be used as a comparative measure of overall hydraulic adaptation across species and to assess the impact of environmental variation, especially drought, on water transport. 2. We summarize the main methods currently in use for measurements of stem xylem hydraulic properties. Measurements can be accomplished in a number of ways, including using a pipette, an analytic balance or a ‘pressure-drop’ flow meter.We provide new details on the design of a relatively inexpensive and easily field-deployable flow meter that is flexible for a variety of applications. The biological challenges associated with these measurements arise from the difficulties of working with diverse living tissues of variable geometry. 3. We provide a review of best practices and provide technical guidance, emphasizing measurements on detached samples using portable equipment.

Increasing drought is one of the most critical challenges facing species and ecosystems worldwide, and improved theory and practices are needed for quantification of species tolerances. Leaf water potential at turgor loss, or wilting (π(tlp) ), is classically recognised as a major physiological determinant of plant water stress response. However, the cellular basis of π(tlp) and its importance for predicting ecological drought tolerance have been controversial. A meta-analysis of 317 species from 72 studies showed that π(tlp) was strongly correlated with water availability within and across biomes, indicating power for anticipating drought responses. We derived new equations giving both π(tlp) and relative water content at turgor loss point (RWC(tlp) ) as explicit functions of osmotic potential at full turgor (π(o) ) and bulk modulus of elasticity (e). Sensitivity analyses and meta-analyses showed that π(o) is the major driver of π(tlp) . In contrast, e plays no direct role in driving drought tolerance within or across species, but sclerophylly and elastic adjustments act to maintain RWC(tlp,) preventing cell dehydration, and additionally protect against nutrient, mechanical and herbivory stresses independent of drought tolerance. These findings clarify biogeographic trends and the underlying basis of drought tolerance parameters with applications in comparative assessments of species and ecosystems worldwide.

 Premise of the study: Hawaii is home to 238 native and 35 alien fern and lycophyte taxa distributed across steep gradients in elevation and resource availability. The fern fl ora spans a wide range of growth forms, with extraordinary diversity in morphology and plant size. Yet the potential factors underlying this diversity have remained enigmatic.  Methods: We used a trait database generated from the most recent and comprehensive survey of Hawaiian ferns and lycophytes to test hypotheses of size-scaling and trait associations with environment and growth form as factors underlying this diversity. We also tested relationships among morphology, taxon abundance and distribution and identifi ed key differences between native and alien taxa.  Key results: Strong trait – trait relationships included geometric scaling of plant dimensions with a tendency for more divided fronds in larger ferns. Trait – environment relationships independent of size included more divided fronds at higher elevation, longer blades in shaded habitats, and fronds with shorter stipes and fewer pinnae in drier habitats. Growth forms differed in mean size with epiphytic and epipetric taxa smaller than terrestrial ferns. Plant size was independent of taxon abundance and distribution across islands, and native and alien ferns did not differ in mean size. Alien taxa were more abundant, especially at lower elevations, apparently due to human land use.  Conclusions: These relationships point to linkages of fern form and demography with biogeography and highlight potential fl ora-scale physiological and morphological adaptations in ferns across contrasting environments.

Summary 1. General hypotheses have related nutrient stoichiometry to physiology and structure for tracheophytes across global resource gradients. These hypotheses have not been tested on bryophytes despite their importance in numerous ecosystem processes. 2. Twelve bryophyte species were sampled across an elevation · soil age matrix representing a dramatic range of climates on Mauna Loa, Hawaii. 3. We tested six major hypotheses for the relationship of tissue nutrients to environment and morphology, and for differences in composition between bryophytes and tracheophytes. 4. Results supported stoichiometric theory. Mass-based nutrient concentrations increased with soil nutrient availability; area-based nutrient concentrations increased with irradiance as mediated by bryophyte canopy mass per area; N and P followed the general scaling shown previously for tracheophytes; and P increased and N:P decreased with elevation consistent with increasing cold tolerance. 5. Synthesis. These findings extend the generality of stoichiometric theory, pointing to convergent physiological responses across distantly related lineages, operating across local and global resource gradients.

Summary 1. Computers are widely used in all aspects of research but their application to in-field data collection for forest plots has rarely been evaluated. 2. We developed digital data collection methods using ESRI mapping software and ruggedized field computers to map and measure 30 000 trees in two 4-ha forest dynamics plots in wet and dry tropical forest in Hawaii. We then compared our data collection and entry effort with published values for other forest dynamics plots with the same tree measurement protocols to estimate the efficiency of our methods relative to the more typical use of paper data collection sheets. 3. In-field data collection effort was comparable for all plots. However, use of digital methods resulted in an average 11AE8% reduction in total effort due to reduced secondary data entry time. 4. The digital data collection methods described in this article can be applied to a wide range of ecological projects, especially long-term research or monitoring projects where mapping can be integrated into data collection.

Summary 1. Leaf structural and physiological traits are associated with growth form and habitat, but little is known of the specific traits associated with hemiepiphytes, which are an important component of many tropical forests. Given their life history that includes a drought prone epiphytic stage, hemiepiphytes should be expected to have more drought tolerance-related traits than co-occurring terrestrial species. 2. The genus Ficus includes woody hemiepiphytes distributed in tropical areas throughout the world. Traits related to the flux of water through the leaf and to drought adaptations were studied in five hemiephiphytic (H) and five non-hemiepiphytic (NH) Ficus tree species grown in a common garden to determine genetically based differences. 3. Leaves of H and NH species differed substantially in structure and physiology; on average, H species had smaller leaves with higher leaf mass per unit area, thicker epidermis, smaller vessel lumen diameters in petioles and lower petiole hydraulic conductivity. Leaf traits also indicated stronger drought tolerance in H species, including lower epidermal conductance and turgor loss point and earlier stomatal closure with desiccation than NH species. Across H and NH species, traits related to water flux capacity were negatively correlated with traits related to drought tolerance. 4. The divergences in hydraulics and water relations between growth forms for these closely related species reflected specialization according to contrasting habitat and life form. Conservative water use and increased ability of leaves to persist under severe drought would provide an advantage for H species, especially during the epiphytic phase, while the higher potential water use of NH species would be associated with higher assimilation rates and competitiveness under high water supply. 5. The results indicate a trade-off between leaf water flux capacity and leaf drought tolerance across these hemiephiphytic and non-hemiepiphytic species. Species adaptation to habitats with contrasting demands on leaf function may lead to divergence along a leaf water-flux-droughttolerance spectrum.

Linkages among leaf traits and environment have most often been tested across communities but infrequently within lineages. We studied seven endemic Hawaiian Plantago taxa radiated across elevations, climates, and habitats. We grew plants of six taxa in controlled conditions for 1–2 yr and collected leaves from a seventh in the field. For all taxa, we measured 46 leaf traits and tested hypotheses for trait-environment and trait-trait associations. Because of the rarity of the study plants, our study included low replication within taxa and multiple growth locations; despite these limitations, given reasonable assumptions, our analyses pointed to genetic differentiation among taxa. The leaves of bog taxa were smaller and thicker than those of woodland taxa, with higher leaf mass per area (LMA), stomatal pore area per leaf area, and carbon isotope discrimination (δ13C). Taxa from higher elevations had thicker leaves and higher LMA, as well as lower nitrogen per mass and higher adaxial stomatal distribution....

Large-fronded tree ferns are critical components of many tropical forests. We investigated frond and whole-plant allometries for Hawaiian keystone species Cibotium glaucum, for prediction and to compare with global scaling relationships. We found that C. glaucum fronds maintain geometric proportionality across a wide range of plant and frond sizes. These relationships result in strong allometries that permit rapid field estimation of frond size from simple linear dimensions. C. glaucum frond allometries complied with intra- and interspecific global trends for leaf area versus mass established for much smaller-leafed species, indicating ‘diminishing returns’ in photosynthetic area per investment in mass for larger fronds. The intraspecific trend was related to declining water content in larger fronds, but not to a significantly larger investment in stipe or rachis relative to lamina. However, C. glaucum complied with the global interspecific trends for greater allocation to support structures in larger leaves. Allometries for frond number and size versus plant height showed that as plants increase in height, frond production and/or retention progressively declines, and the increases of leaf size tend to level off. These frond and whole plant-level relationships indicate the potential for estimating frond area and mass at landscape scale to enrich studies of forest dynamics.

Summary 1Populations of Metrosideros polymorpha establish across a broad range of precipitation in Hawai‘i – from 10000 mm per year. To determine whether adjustment of hydraulic and photosynthetic traits could contribute to this success in both high and low rainfall, we sampled populations on the wet and the dry sides of Hawai‘i Island, replicated on two different-aged lava flows at similar elevation and mean annual temperature. 2We quantified 24 stem and leaf traits in an integrated study of hydraulics, gas exchange, leaf chemistry and anatomy. 3Values for traits associated with capacity for water transport through the plant and gas exchange per leaf area were higher at dry sites, including photosynthetic capacity, nitrogen per leaf area and hydraulic conductivity on both sapwood area and leaf area basis. These adjustments, due to plastic and/or heritable differentiation, would partially compensate for generally lower water availability. 4Specific leaf area shifted towards lower values at the dry sites. However, several other traits associated directly with drought tolerance were inflexible across sites, including stem vulnerability to embolism and leaf cuticular conductance, indicating that the ability to persist through dry periods is sustained across the species’ range. 5The ability of M. polymorpha to establish across a wide range of habitats is associated with, on one hand, adjustments in traits that would enable sustained growth across a dramatic range of moisture supply while simultaneously maintaining as fixed other traits that would contribute to survival through extended drought.

• Premise of the study: The diverse early Eocene flora from Laguna del Hunco (LH) in Patagonia, Argentina has many nearest living relatives (NLRs) in Australasia but few in South America, indicating the differential survival of an ancient, trans‐Antarctic rainforest biome. To better understand this significant biogeographic pattern, we used detailed comparisons of leaf size and floristics to quantify the legacy of LH across a large network of Australian rainforest‐plot assemblages. • Methods: We applied vein scaling, a new method for estimating the original areas of fragmented leaves. We then compared leaf size and floristics at LH with living Australian assemblages and tabulated the climates of those where NLRs occur, along with additional data on climatic ranges of “ex‐Australian” NLRs that survive outside of Australia. • Key results: Vein scaling estimated areas as accurately as leaf‐size classes. Applying vein scaling to fossil fragments increased the grand mean area of LH by 450 mm2, recovering more originally large leaves. The paleoflora has a majority of microphyll leaves, comparable to leaf litter in subtropical Australian forests, which also have the greatest floristic similarity to LH. Tropical Australian assemblages also share many taxa with LH, and ex‐Australian NLRs mostly inhabit cool, wet montane habitats no longer present in Australia. • Conclusions: Vein scaling is valuable for improving the resolution of fossil leaf‐size distributions by including fragmented specimens. The legacy of LH is evident not only in subtropical and tropical Australia but also in tropical montane Australasia and Southeast Asia, reflecting the disparate histories of surviving Gondwanan lineages. Fil: Merkhofer, Lisa. State University of Pennsylvania; Estados Unidos Fil: Wilf, Peter. State University of Pennsylvania; Estados Unidos Fil: Haas, M. Tyler. State University of Pennsylvania; Estados Unidos Fil: Kooyman, Robert M.. Macquarie University; Australia Fil: Sack, Lawren. University of California at Los Angeles; Estados Unidos Fil: Scoffoni, Christine. University of California at Los Angeles; Estados Unidos Fil: Cúneo, Néstor Rubén. Museo Paleontológico Egidio Feruglio; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina

15 páginas.-- 6 figuras.-- 6 tablas.-- 60 referencias.-- Supplementary data related to this article can be found at http://dx.doi.org/10.1016/j.jaridenv.2015.03.001. or http://www.sciencedirect.com/science/article/pii/S0140196315000671#ec1, © 2015 Elsevier Ltd. Much attention has been paid to differences in leaf form and composition among vegetation types, but less to the frequently substantial variation within vegetation types. We focused on the extent to which correlations between variables are the same in both succulent-poor and succulent-rich vegetation in semi-arid SE Spain. Mean foliar [N] of perennials varied among species over a 5-fold range. Across species, [N] was positively correlated with specific leaf area (i.e., leaf area divided by dry mass; SLA) and with water concentration at saturation (WCS) in the grasslands, excluding the one succulent species. In succulent-rich vegetation on marl, SLA was correlated with [N] but not WCS, and there was a wedge-shaped relationship between [N] and WCS. Foliar [N] and [P] were positively correlated in the grasslands, but not in succulent-rich vegetation on marl. The N/P quotient varied from 8 to 29, with mean 14 in grassland on limestone and mean 26 in grassland on deep soil over gypsum. Our chief finding is that most correlations among SLA, WCS, [N] and [P] found in the non-succulent vegetation are not found in the succulent-rich vegetation. The results are discussed in relation to global patterns and the problems of defining succulence., Thank the Consejería de Medio Ambiente, Junta de Andalucía for permission to collect plant samples. Glyn Jones at the University of Cambridge did the chemical analyses of leaves; the Servicio de Análisis at IRNAS, CSIC, Sevilla analyzed the whole shoots and soil samples, thanks to L.V. García for his help and advice. Funds to make this study were provided by the Spanish Ministry of Science (projects PB97-1177, CGL2010-17081 and DiverbosCGL2011-30285-C02) and European FEDER.

Species variation in functional traits may reflect diversification relating to convergence and/or divergence depending on environmental pressures and phylogenetic history. We tested trait-environment relationships and their basis in finer-scale evolutionary processes among nine extant Hawaiian species of Scaevola L. (Goodeniaceae), a taxon with a complex history of three independent colonizations by different phylogenetic lineages, parallel ecological specialization, and homoploid hybridization events in Hawai'i. Using a wild population for each species, we evaluated traits related to plant function (morphology, leaf and wood anatomy, nutrient and carbon isotope composition). Hawaiian Scaevola species were distributed across coastal, dry forest and wet forest environments; multivariate environmental analysis using abiotic and biotic factors further showed that species from distantly related lineages inhabited similar environments. Many traits correlated with environment (based on the multivariate environmental analysis), considering both distantly related species and more closely related species. Scaevola species within shared habitats generally showed trait convergence across distantly related lineages, particularly among wet forest species. Furthermore, trait diversification through divergence was extensive among closely related Scaevola species that radiated into novel environments, especially in plant morphology and traits affecting water relations. Homoploid hybrid-origin species were "intermediate" compared to their ancestral parent species, and possessed trait combinations relevant for their current habitat. The diversity in functional traits reflected strong influences of both ecology and evolutionary history in native Hawaiian Scaevola species, and trait correspondence with environment was due to the combination of multiple processes within the taxon: trait pre-adaptation and filtering, evolutionary convergence, divergence, and hybridization.