Continued adaptation of C4 photosynthesis after an initial burst of changes in the Andropogoneae grasses

C4 photosynthesis is a complex trait that sustains fast growth and high productivity in tropical and subtropical conditions and evolved repeatedly in flowering plants. One of the major C4 lineages is Andropogoneae, a group of ~ 1,200 grass species that includes some of the world's most important crops and species dominating tropical and some temperate grasslands. Previous efforts to understand C4 evolution in the group have compared a few model C4 plants to distantly related C3 species, so that changes directly responsible for the transition to C4 could not be distinguished from those that preceded or followed it. In this study, we analyse the genomes of 66 grass species, capturing the earliest diversification within Andropogoneae as well as their C3 relatives. Phylogenomics combined with molecular dating and analyses of protein evolution show that many changes linked to the evolution of C4 photosynthesis in Andropogoneae happened in the Early Miocene, between 21 and 18 Ma, after the sp...