Multiple lines of evidence suggest that plant water-use efficiency (WUE)—the ratio of carbon assimilation to water loss—has increased in recent decades. Although rising atmospheric CO ₂ has been proposed as the principal cause, the underlying physiological mechanisms are still being debated, and implications for the global water cycle remain uncertain. Here, we addressed this gap using 30-y tree ring records of carbon and oxygen isotope measurements and basal area increment from 12 species in 8 North American mature temperate forests. Our goal was to separate the contributions of enhanced photosynthesis and reduced stomatal conductance to WUE trends and to assess consistency between multiple commonly used methods for estimating WUE. Our results show that tree ring-derived estimates of increases in WUE are consistent with estimates from atmospheric measurements and predictions based on an optimal balancing of carbon gains and water costs, but are lower than those based on ecosystem-scale flux observations. Although both physiological mechanisms contributed to rising WUE, enhanced photosynthesis was widespread, while reductions in stomatal conductance were modest and restricted to species that experienced moisture limitations. This finding challenges the hypothesis that rising WUE in forests is primarily the result of widespread, CO ₂ -induced reductions in stomatal conductance.

Sampling of 1367 trees was conducted in the Side wildfire (4 May 1996), Bridger-Knoll wildfire (20 June 1996) and Dauber prescribed fire (9 September 1995) in northern Arizona ponderosa pine forests ( Pinus ponderosa ). Tree mortality was assessed for 3 years after each fire. Three-year post-fire mortality was 32.4% in the Side wildfire, 18.0% in the Dauber prescribed fire, and 13.9% in the Bridger-Knoll wildfire. In the Dauber and Side fires, 95% and 94% of 3-year post-fire mortality occurred by year 2, versus 76% in the Bridger-Knoll wildfire. Compared with trees that lived for 3 years after fire, dead trees in all fires had more crown scorch, crown consumption, bole scorch, ground char, and bark beetle attacks. Logistic regression models were used to provide insight on factors associated with tree mortality after fire. A model using total crown damage by fire (scorch + consumption) and bole char severity as independent variables was the best two-variable model for predicting individual tree mortality for all fires. The amount of total crown damage associated with the onset of tree mortality decreased as bole char severity increased. Models using diameter at breast height (dbh) and crown volume damage suggested that tree mortality decreased as dbh increased in the Dauber prescribed fire where trees were smallest, and tree mortality increased as dbh increased in the Side and Bridger-Knoll wildfires where trees were largest. Moreover, a U-shaped dbh–mortality distribution for all fires suggested higher mortality for the smallest and largest trees compared with intermediate-size trees. We concluded that tree mortality is strongly influenced by interaction between crown damage and bole char severity, and differences in resistance to fire among different-sized trees can vary among sites.

Despite their sparse vegetation, dryland regions exert a huge influence over global biogeochemical cycles because they cover more than 40% of the world surface (Schimel 2010 Science 327 418–9). It is thought that drylands dominate the inter-annual variability (IAV) and long-term trend in the global carbon (C) cycle (Poulter et al 2014 Nature 509 600–3, Ahlstrom et al 2015 Science 348 895–9, Zhang et al 2018 Glob. Change Biol. 24 3954–68). Projections of the global land C sink therefore rely on accurate representation of dryland C cycle processes; however, the dynamic global vegetation models (DGVMs) used in future projections have rarely been evaluated against dryland C flux data. Here, we carried out an evaluation of 14 DGVMs (TRENDY v7) against net ecosystem exchange (NEE) data from 12 dryland flux sites in the southwestern US encompassing a range of ecosystem types (forests, shrub- and grasslands). We find that all the models underestimate both mean annual C uptake/release as well as the magnitude of NEE IAV, suggesting that improvements in representing dryland regions may improve global C cycle projections. Across all models, the sensitivity and timing of ecosystem C uptake to plant available moisture was at fault. Spring biases in gross primary production (GPP) dominate the underestimate of mean annual NEE, whereas models’ lack of GPP response to water availability in both spring and summer monsoon are responsible for inability to capture NEE IAV. Errors in GPP moisture sensitivity at high elevation forested sites were more prominent during the spring, while errors at the low elevation shrub and grass-dominated sites were more important during the monsoon. We propose a range of hypotheses for why model GPP does not respond sufficiently to changing water availability that can serve as a guide for future dryland DGVM developments. Our analysis suggests that improvements in modeling C cycle processes across more than a quarter of the Earth’s land surface could be achieved by addressing the moisture sensitivity of dryland C uptake.