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.