One of the most striking ecological trends is the association of small leaves with dry and cold climates, described 2400 years ago by Theophrastus, and recently recognized for eudicotyledonous plants at the global scale 1-3 . For eudicotyledons, this pattern is attributed 24 to small leaves having a thinner boundary layer to avoid extreme leaf temperatures 4 , and 25 their developing vein traits that improve water transport under cold or dry climates 5,6 . Yet, 26 the global distribution of leaf size and its mechanisms have not been tested in grasses, an 27 extraordinarily diverse lineage, distinct in leaf morphology, which contributes 33% of 28 terrestrial primary productivity, including the bulk of crop production 7 . Here we demonstrate that grasses have shorter and narrower leaves under colder and drier climates worldwide. We show that small grass leaves have thermal advantages and vein development that contrast with those of eudicotyledons, but that also explain the abundance of small leaves in cold and dry climates. The worldwide distribution of grass leaf size exemplifies how biophysical and developmental processes result in convergence across major lineages in adaptation to climate globally, and highlights the importance of leaf size and venation architecture for grass performance in past, present and future ecosystems. Data Fig. 1, SupplementaryTable 3). We tested whether developmental scaling would confer 64 small leaves with potential climatic advantages. 65 4 66 Box 1. Synthetic model of grass leaf vein development based on published data for 20 species (Supplementary Tables 5-6), conferring small leaves with traits advantageous under cold and dry climates Grass leaf development includes five phases based on developmental zones: Phase P (formation and expansion of the primordium, P): "Founder cells" in the periphery of the shoot apical meristem generate the leaf primordium. Cell divisions drive growth of a hood-like structure, in which the central 1° vein (midvein) and the large 2° veins are initiated early and extend acropetally, enabling their prolonged diameter growth (Box 1 Fig. 1a, c, e). Henceforth, discrete spatial growth zones develop at the leaf base and drive leaf expansion laterally and longitudinally.
Phase D (formation of the cell division zone, DZ):The basal cell division zone (DZ) expands slightly, driving minimal growth (Box 1 Fig. 1a, b). The 1° and 2° vein orders (major veins) complete their patterning basipetally along the leaf blade and increase in diameter (Box 1 Fig. 1c, e). Meanwhile, beginning at the lamina tip, C 3 species form a single order of small longitudinal minor veins, i.e., 3° veins, as do most C 4 species, i.e., C 4-3L species. Some C 4 species of the subfamily Panicoideae additionally form smaller 4° veins, i.e., C 4-4L species 15 (Box 1 Fig. 1c).
Phase D-E (DZ, and formation of the expansion zone, EZ):Cells from the DZ transition to a distinct, distal expansion zone (EZ).In the EZ, cell expansion in width and length spaces apart the 1° and 2° veins, resulting in th...