• adaptive dynamic global vegetation model (aDGVM);
  • C4 expansion;
  • CO2 starvation hypothesis;
  • dynamic vegetation model;
  • fire adaptation;
  • grassland;
  • late Miocene;
  • savanna


  • Large proportions of the Earth’s land surface are covered by biomes dominated by C4 grasses. These C4-dominated biomes originated during the late Miocene, 3–8 million years ago (Ma), but there is evidence that C4 grasses evolved some 20 Ma earlier during the early Miocene/Oligocene. Explanations for this lag between evolution and expansion invoke changes in atmospheric CO2, seasonality of climate and fire. However, there is still no consensus about which of these factors triggered C4 grassland expansion.
  • We use a vegetation model, the adaptive dynamic global vegetation model (aDGVM), to test how CO2, temperature, precipitation, fire and the tolerance of vegetation to fire influence C4 grassland expansion. Simulations are forced with late Miocene climates generated with the Hadley Centre coupled ocean–atmosphere–vegetation general circulation model.
  • We show that physiological differences between the C3 and C4 photosynthetic pathways cannot explain C4 grass invasion into forests, but that fire is a crucial driver. Fire-promoting plant traits serve to expand the climate space in which C4-dominated biomes can persist.
  • We propose that three mechanisms were involved in C4 expansion: the physiological advantage of C4 grasses under low atmospheric CO2 allowed them to invade C3 grasslands; fire allowed grasses to invade forests; and the evolution of fire-resistant savanna trees expanded the climate space that savannas can invade.