Global convergence in the balance between leaf water supply and demand across vascular land plants

Coordination between the density of veins (water supply) and stomata (demand for water) has been found in the leaves of modern angiosperms and also in ferns. This suggests that this coordinated development is not a unique adaptation of derived angiosperms that enables their high productivity. To test this, we compiled leaf vein and stomatal density data from 520 land vascular plant species including derived angiosperms, basal angiosperms, gymnosperms and ferns. We found global coordination across vascular land plants, although the relationships were not significant in gymnosperms and vessel-less angiosperms. By comparing the evolution of xylem conduit elements with variation in the density of veins and stomata and theoretical stomatal conductance among plant lineages, we found that the physiological advantage of modern angiosperms is associated with the emergence of xylem with low intraconduit resistance and leaves with high vein and stomata densities. Thus our results indicate two major events associated with surges in xylem hydraulic capacity in angiosperms: (1) the origin of vessels and (2) the emergence of vessels with simple perforation plates, which diminished physical limitations on stomatal conductance. These evolutionary innovations may have enabled derived angiosperms to be more productive and adaptive to the changing climate.