Mediterranean-type ecosystems (MTEs) possess the highest levels of plant species richness in the world outside of the wet tropics. Sclerophyll vegetation similar to today’s mediterranean-type shrublands was already present on oligotrophic soils in the wet and humid climate of the Cretaceous, with fire-adapted Paleogene lineages in southwestern Australia and the Cape Region. The novel MTC seasonality present since the mid-Miocene has allowed colonization of MTEs from a regional species pool with associated diversification. Fire persistence has been a primary driving factor for speciation in four of the five regions. Understanding the regional patterns of plant species diversity among the MTEs involves complex interactions of geologic and climatic histories for each region as well as ecological factors that have promoted diversification in the Neogene and Quaternary. A critical element of species richness for many MTE lineages has been their ability to speciate and persist at fine spatial scales, with low rates of extinction.