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            Abstract PremisePrevious studies have suggested a trade‐off between trichome density (Dt) and stomatal density (Ds) due to shared cell precursors. We clarified how, when, and why this developmental trade‐off may be overcome across species. MethodsWe derived equations to determine the developmental basis forDtandDsin trichome and stomatal indices (itandis) and the sizes of epidermal pavement cells (e), trichome bases (t), and stomata (s) and quantified the importance of these determinants ofDtandDsfor 78 California species. We compiled 17 previous studies ofDt–Dsrelationships to determine the commonness ofDt–Dsassociations. We modeled the consequences of differentDt–Dsassociations for plant carbon balance. ResultsOur analyses showed that higherDtwas determined by higheritand lowere, and higherDsby higherisand lowere. Across California species, positiveDt–Dscoordination arose due toit–iscoordination and impacts of the variation ine. ADt–Dstrade‐off was found in only 30% of studies. Heuristic modeling showed that species sets would have the highest carbon balance with a positive or negative relationship or decoupling ofDtandDs, depending on environmental conditions. ConclusionsShared precursor cells of trichomes and stomata do not limit higher numbers of both cell types or drive a generalDt–Dstrade‐off across species. This developmental flexibility across diverse species enables differentDt–Dsassociations according to environmental pressures. Developmental trait analysis can clarify how contrasting trait associations would arise within and across species.more » « less
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            Abstract The relationship between stomatal traits and environmental drivers across plant communities has important implications for ecosystem carbon and water fluxes, but it has remained unclear. Here, we measure the stomatal morphology of 4492 species-site combinations in 340 vegetation plots across China and calculate their community-weighted values for mean, variance, skewness, and kurtosis. We demonstrate a trade-off between stomatal density and size at the community level. The community-weighted mean and variance of stomatal density are mainly associated with precipitation, while that of stomatal size is mainly associated with temperature, and the skewness and kurtosis of stomatal traits are less related to climatic and soil variables. Beyond mean climate variables, stomatal trait moments also vary with climatic seasonality and extreme conditions. Our findings extend the knowledge of stomatal trait–environment relationships to the ecosystem scale, with applications in predicting future water and carbon cycles.more » « less
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            Summary A surge of papers have reported low leaf vulnerability to xylem embolism during drought. Here, we focus on the less studied, and more sensitive, outside‐xylem leaf hydraulic responses to multiple internal and external conditions. Studies of 34 species have resolved substantial vulnerability to dehydration of the outside‐xylem pathways, and studies of leaf hydraulic responses to light also implicate dynamic outside‐xylem responses. Detailed experiments suggest these dynamic responses arise at least in part from strong control of radial water movement across the vein bundle sheath. While leaf xylem vulnerability may influence leaf and plant survival during extreme drought, outside‐xylem dynamic responses are important for the control and resilience of water transport and leaf water status for gas exchange and growth.more » « less
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            Abstract Tree diversity can promote both predator abundance and diversity. However, whether this translates into increased predation and top‐down control of herbivores across predator taxonomic groups and contrasting environmental conditions remains unresolved. We used a global network of tree diversity experiments (TreeDivNet) spread across three continents and three biomes to test the effects of tree species richness on predation across varying climatic conditions of temperature and precipitation. We recorded bird and arthropod predation attempts on plasticine caterpillars in monocultures and tree species mixtures. Both tree species richness and temperature increased predation by birds but not by arthropods. Furthermore, the effects of tree species richness on predation were consistent across the studied climatic gradient. Our findings provide evidence that tree diversity strengthens top‐down control of insect herbivores by birds, underscoring the need to implement conservation strategies that safeguard tree diversity to sustain ecosystem services provided by natural enemies in forests.more » « less
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            Abstract Allometric relationships among the dimensions of leaves and their cells hold across diverse eudicotyledons, but have remained untested in the leaves of grasses. We hypothesised that geometric (proportional) allometries of cell sizes across tissues and of leaf dimensions would arise due to the coordination of cell development and that of cell functions such as water, nutrient and energy transport, and that cell sizes across tissues would be associated with light‐saturated photosynthetic rate. We tested predictions across 27 globally distributed C3and C4grass species grown in a common garden. We found positive relationships among average cell sizes within and across tissues, and of cell sizes with leaf dimensions. Grass leaf anatomical allometries were similar to those of eudicots, with exceptions consistent with the fewer cell layers and narrower form of grass leaves, and the specialised roles of epidermis and bundle sheath in storage and leaf movement. Across species, mean cell sizes in each tissue were associated with light‐saturated photosynthetic rate per leaf mass, supporting the functional coordination of cell sizes. These findings highlight the generality of evolutionary allometries within the grass lineage and their interlinkage with coordinated development and function.more » « less
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            Abstract Intra‐specific trait variation (ITV) plays a role in processes at a wide range of scales from organs to ecosystems across climate gradients. Yet, ITV remains rarely quantified for many ecophysiological traits typically assessed for species means, such as pressure volume (PV) curve parameters including osmotic potential at full turgor and modulus of elasticity, which are important in plant water relations. We defined a baseline “reference ITV” (ITVref) as the variation among fully exposed, mature sun leaves of replicate individuals of a given species grown in similar, well‐watered conditions, representing the conservative sampling design commonly used for species‐level ecophysiological traits. We hypothesized that PV parameters would show low ITVrefrelative to other leaf morphological traits, and that their intraspecific relationships would be similar to those previously established across species and proposed to arise from biophysical constraints. In a database of novel and published PV curves and additional leaf structural traits for 50 diverse species, we found low ITVreffor PV parameters relative to other morphological traits, and strong intraspecific relationships among PV traits. Simulation modeling showed that conservative ITVrefenables the use of species‐mean PV parameters for scaling up from spectroscopic measurements of leaf water content to enable sensing of leaf water potential.more » « less
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            Abstract Improved estimation of climate niches is critical, given climate change. Plant adaptation to climate depends on their physiological traits and their distributions, yet traits are rarely used to inform the estimation of species climate niches, and the power of a trait‐based approach has been controversial, given the many ecological factors and methodological issues that may result in decoupling of species' traits from their native climate.For 107 species across six ecosystems of California, we tested the hypothesis that mechanistic leaf and wood traits can robustly predict the mean of diverse species' climate distributions, when combining methodological improvements from previous studies, including standard trait measurements and sampling plants growing together at few sites. Further, we introduce an approach to quantify species' trait‐climate mismatch.We demonstrate a strong power to predict species mean climate from traits. As hypothesized, the prediction of species mean climate is stronger (and mismatch lower) when traits are sampled for individuals closer to species' mean climates.Improved resolution of species' climate niches based on mechanistic traits can importantly inform conservation of vulnerable species under the threat of climatic shifts in upcoming decades. Read the freePlain Language Summaryfor this article on the Journal blog.more » « less
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            Abstract The diversity of specialized molecules produced by plants radiating along ecological gradients is thought to arise from plants' adaptations to local conditions. Therefore, closely related species growing in similar habitats should phylogenetically converge, or diverge, in response to similar climates, or similar interacting animal communities. We here asked whether closely related species in the genusHaplopappus(Asteraceae) growing within the same elevation bands in the Andes, converged to produce similar floral odors. To do so, we combine untargeted analysis of floral volatile organic compounds with insect olfactory bioassay in congenericHaplopappus(Asteraceae) species growing within the same elevation bands along the Andean elevational gradient. We then asked whether the outcome of biotic interactions (i.e., pollination vs. seed predation) would also converge across species within the same elevation. We found that flower odors grouped according to their elevational band and that the main floral visitor preferred floral heads from low‐elevation band species. Furthermore, the cost–benefit ratio of predated versus fertilized seeds was consistent within elevation bands, but increased with elevation, from 6:1 at low to 8:1 at high elevations. In the light of our findings, we propose that climate and insect community changes along elevation molded a common floral odor blend, best adapted for the local conditions. Moreover, we suggest that at low elevation where floral resources are abundant, the per capita cost of attracting seed predators is diluted, while at high elevation, sparse plants incur a higher herbivory cost per capita. Together, our results suggest that phytochemical convergence may be an important factor driving plant–insect interactions and their ecological outcomes along ecological gradients.more » « less
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            Abstract Frequent drought and high temperature conditions in California vineyards necessitate plant stress detection to support irrigation management strategies and decision making. Remote sensing provides a powerful tool to continuously monitor vegetation function across spatial and temporal scales. In this study, we utilized a tower-based optical-remote sensing system to continuously monitor four vineyard subplots in California’s Central Valley. We compared the performance of the greenness-based normalized difference vegetation index (NDVI) and the physiology-based photochemical reflectance index (PRI) to track variations of eddy covariance estimated gross primary productivity (GPP) during four stress events between July and September 2020. Our results demonstrate that NDVI was invariant during stress events. In contrast, PRI was effective at tracking the short-term stress-induced declines and recovery of GPP associated with soil water depletion and increased air temperature, as well as reductions in GPP from decreased PAR caused by smokey conditions from nearby fires. Canopy-scale remote sensing can provide continuous real-time data, and physiology-based vegetation indices such as PRI can be used to monitor variation of photosynthetic activity during stress events to aid in management decisions.more » « less
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            Summary Rising temperatures are influencing forests on many scales, with potentially strong variation vertically across forest strata. Using published research and new analyses, we evaluate how microclimate and leaf temperatures, traits, and gas exchange vary vertically in forests, shaping tree, and ecosystem ecology. In closed‐canopy forests, upper canopy leaves are exposed to the highest solar radiation and evaporative demand, which can elevate leaf temperature (Tleaf), particularly when transpirational cooling is curtailed by limited stomatal conductance. However, foliar traits also vary across height or light gradients, partially mitigating and protecting against the elevation of upper canopyTleaf. Leaf metabolism generally increases with height across the vertical gradient, yet differences in thermal sensitivity across the gradient appear modest. Scaling from leaves to trees, canopy trees have higher absolute metabolic capacity and growth, yet are more vulnerable to drought and damagingTleafthan their smaller counterparts, particularly under climate change. By contrast, understory trees experience fewer extreme highTleaf's but have fewer cooling mechanisms and thus may be strongly impacted by warming under some conditions, particularly when exposed to a harsher microenvironment through canopy disturbance. As the climate changes, integrating the patterns and mechanisms reviewed here into models will be critical to forecasting forest–climate feedback.more » « less
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