Plant ecological strategies are shaped by numerous functional traits and their trade‐offs. Trait network analysis enables testing hypotheses for the shifting of trait correlation architecture across communities differing in climate and productivity. We built plant trait networks (PTNs) for 118 species within six communities across an aridity gradient, from forest to semi‐desert across the California Floristic Province, based on 34 leaf and wood functional traits, representing hydraulic and photosynthetic function, structure, economics and size. We developed hypotheses for the association of PTN parameters with climate and ecosystem properties, based on theory for the adaptation of species to low resource/stressful environments versus higher resource availability environments with greater potential niche differentiation. Thus, we hypothesized that across community PTNs, trait network connectivity (i.e., the degree that traits are intercorrelated) and network complexity (i.e., the number of trait modules, and the degree of trait integration among modules) would be lower for communities adapted to arid climates and higher for communities adapted to greater water availability, similarly to trends expected for phylogenetic diversity, functional richness and productivity. Further, within given PTNs, we hypothesized that traits would vary strongly in their network connectivity and that the traits most centrally connected within PTNs would be those with the least across‐species variation. Across communities from more arid to wetter climates, PTN architecture varied from less to more interconnected and complex, in association with functional richness, but PTN architecture was independent of phylogenetic diversity and ecosystem productivity. Within the community PTNs, traits with lower species variation were more interconnected. Synthesis. The responsiveness of PTN architecture to climate highlights how a wide range of traits contributes to physiological and ecological strategies with an architecture that varies among plant communities. Communities in more arid environments show a lower degree of phenotypic integration, consistent with lesser niche differentiation. Our study extends the usefulness of PTNs as an approach to quantify tradeoffs among multiple traits, providing connectivity and complexity parameters as tools that clarify plant environmental adaptation and patterns of trait associations that would influence species distributions, community assembly, and ecosystem resilience in response to climate change.

Leaf surface conductance to water vapor and CO2 across the epidermis (gleaf) strongly determines the rates of gas exchange. Thus, clarifying the drivers of gleaf has important implications for resolving the mechanisms of photosynthetic productivity and leaf and plant responses and tolerance to drought. It is well recognized that gleaf is a function of the conductances of the stomata (gs) and of the epidermis + cuticle (gec). Yet, controversies have arisen around the relative roles of stomatal density (d) and size (s), fractional stomatal opening (α; aperture relative to maximum), and gec in determining gleaf. Resolving the importance of these drivers is critical across the range of leaf surface conductances, from strong stomatal closure under drought (gleaf,min), to typical opening for photosynthesis (gleaf,op), to maximum achievable opening (gleaf,max). We derived equations and analyzed a compiled database of published and measured data for approximately 200 species and genotypes. On average, within and across species, higher gleaf,min was determined 10 times more strongly by α and gec than by d and negligibly by s; higher gleaf,op was determined approximately equally by α (47%) and by stomatal anatomy (45% by d and 8% by s), and negligibly by gec; and higher gleaf,max was determined entirely by d. These findings clarify how diversity in stomatal functioning arises from multiple structural and physiological causes with importance shifting with context. The rising importance of d relative to α, from gleaf,min to gleaf,op, enables even species with low gleaf,min, which can retain leaves through drought, to possess high d and thereby achieve rapid gas exchange in periods of high water availability.

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 free Plain Language Summary for this article on the Journal blog.

Premise: Previous studies have suggested a trade‐off between trichome density (D_t) and stomatal density (D_s) due to shared cell precursors. We clarified how, when, and why this developmental trade‐off may be overcome across species.Methods: We derived equations to determine the developmental basis for D_t and D_s in trichome and stomatal indices (i_t and i_s) and the sizes of epidermal pavement cells (e), trichome bases (t), and stomata (s) and quantified the importance of these determinants of D_t and D_s for 78 California species. We compiled 17 previous studies of D_t–D_s relationships to determine the commonness of D_t–D_s associations. We modeled the consequences of different D_t–D_s associations for plant carbon balance.Results: Our analyses showed that higher D_t was determined by higher i_t and lower e, and higher D_s by higher i_s and lower e. Across California species, positive D_t–D_s coordination arose due to i_t–i_s coordination and impacts of the variation in e. A D_t–D_s trade‐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 of D_t and D_s, depending on environmental conditions.Conclusions: Shared precursor cells of trichomes and stomata do not limit higher numbers of both cell types or drive a general D_t–D_s trade‐off across species. This developmental flexibility across diverse species enables different D_t–D_s associations according to environmental pressures. Developmental trait analysis can clarify how contrasting trait associations would arise within and across species.