EVIDENCE FOR ADAPTATION TO FIRE REGIMES IN THE TROPICAL SAVANNAS OF THE BRAZILIAN CERRADO

A recent controversy concerns whether plant traits that are assumed to be adaptations to fire originally evolved in response to selective factors other than fire. We contribute to this debate by investigating the evolution of the endemic woody flora of the fire-prone Cerrado of central Brazil, the most species-rich savanna in the world. We review evidence from dated phylogenies and show that Cerrado lineages started to diversify less than 10 million years ago. These Cerrado lineages are characterized by fire-resistant traits such as thick, corky bark and root sprouting, which have been considered to have evolved as adaptations to drought or nutrient-deficient soils. However, the fact that the lineages carrying these features arose coincident with the rise to dominance of flammable C-4 grasses and expansion of the savanna biome worldwide, and postdating the earlier origin of seasonal climates and the nutrient-poor, acid Cerrado soils suggests that such traits should be considered as adaptations to fire regimes. The nature of these features as adaptations to fire is further suggested by their absence or poor development in related lineages found in fire-free environments with similar edaphic conditions to the Cerrado and by their repeated independent origin in diverse lineages. We present evidence to demonstrate that the evolutionary barrier to entry to the Cerrado is a weak one, presumably because of the ease of evolution of the necessary adaptations to fire regimes for lineages inhabiting neighboring fire-free biomes.