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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

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.

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.

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.

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.

It is often assumed that if a plant species has a higher relative growth rate (RGR) than another species in deep shade, it will have a lower RGR at high irradiance (Spurr & Barnes 1980; Thomas & Bazzaz 1999; Walter 1973). In other words, species change rank (crossover) between low and high irradiance. This idea was suggested chiefly by the finding that in deep shade the massbased net photosynthetic rates of the leaves of shade plants exceed those of sun plants, while at high irradiance the reverse is true (Bjdrkman & Holmgren 1963; Boardman 1977; Givnish 1988). This finding has been assumed to scale up to the level of whole-plant RGR (Shugart 1984). In the past decade, however, a contrary view concerning RGRs has emerged, the idea that if a plant grows faster than another at high irradiance it will also do so in the shade (Kitajima 1994; Poorter 1999). According to this later view, light plays a role in maintaining the mixture of forest species only through the well established trade-off between survival rate in deep shade and RGR in bright light (Kitajima 1994, 1996). Similar experimental studies of RGR responses to irradiance for woody seedlings report surprisingly different results. Here we indicate why such disparate results have been produced. We provide a simple analytical approach to understanding why crossovers should occur among particular species at particular stages of ontogeny. This approach is useful for understanding the maintenance of forest species richness, as well as for interpreting plant specialization in physiology and morphology to contrasting irradiance regimes.