Stomatal and non-stomatal limitations in savanna trees and C4 grasses grown at low, ambient and high atmospheric CO2

By the end of the century, atmospheric CO2 concentration ([CO2]a) could reach 800 ppm, having risen from ~200 ppm ~24 Myr ago. Carbon dioxide enters plant leaves through stomata that limit CO2 diffusion and assimilation, imposing stomatal limitation (LS). Other factors limiting assimilation are collectively called non- stomatal limitations (LNS). C4 photosynthesis concentrates CO2 around Rubisco, typically reducing LS. C4- dominated savanna grasslands expanded under low [CO2]a and are metastable ecosystems where the response of trees and C4 grasses to rising [CO2]a will determine shifting vegetation patterns. How LS and LNS differ between savanna trees and C4 grasses under different [CO2]a will govern the responses of CO2 fixation and plant cover to [CO2]a - but quantitative comparisons are lacking. We measured assimilation, within soil wetting-drying cycles, of three C3 trees and three C4 grasses grown at 200, 400 or 800 ppm [CO2]a. Using assimilation-response curves, we resolved LS and LNS and show that rising [CO2]a alleviated LS, particularly for the C3 trees, but LNS was unaffected and remained substantially higher for the grasses across all [CO2]a treatments. Because LNS incurs higher metabolic costs and recovery compared with LS, our findings indicate that C4 grasses will be compara- tively disadvantaged as [CO2]a rises.