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The dominant vegetation over much of the global land surface is not predetermined by contemporary climate, but also influenced by past environmental conditions. This confounds attempts to predict current and future biome distributions, because even a perfect model would project multiple possible biomes without knowledge of the historical vegetation state. Here we compare the distribution of tree- and grass-dominated biomes across Africa simulated using a dynamic global vegetation model (DGVM). We explicitly evaluate where and under what conditions multiple stable biome states are possible for current and projected future climates. Our simulation results show that multiple stable biomes states are possible for vast areas of tropical and subtropical Africa under current conditions. Widespread loss of the potential for multiple stable biomes states is projected in the 21st Century, driven by increasing atmospheric CO2 . Many sites where currently both tree-dominated and grass-dominated biomes are possible become deterministically tree-dominated. Regions with multiple stable biome states are widespread and require consideration when attempting to predict future vegetation changes. Testing for behaviour characteristic of systems with multiple stable equilibria, such as hysteresis and dependence on historical conditions, and the resulting uncertainty in simulated vegetation, will lead to improved projections of global change impacts., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

Large proportions of the Earth's land surface are covered by biomes dominated by C(4) grasses. These C(4)-dominated biomes originated during the late Miocene, 3-8 million years ago (Ma), but there is evidence that C(4) grasses evolved some 20 Ma earlier during the early Miocene/Oligocene. Explanations for this lag between evolution and expansion invoke changes in atmospheric CO(2), seasonality of climate and fire. However, there is still no consensus about which of these factors triggered C(4) grassland expansion. We use a vegetation model, the adaptive dynamic global vegetation model (aDGVM), to test how CO(2), temperature, precipitation, fire and the tolerance of vegetation to fire influence C(4) 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 C(3) and C(4) photosynthetic pathways cannot explain C(4) grass invasion into forests, but that fire is a crucial driver. Fire-promoting plant traits serve to expand the climate space in which C(4)-dominated biomes can persist. We propose that three mechanisms were involved in C(4) expansion: the physiological advantage of C(4) grasses under low atmospheric CO(2) allowed them to invade C(3) grasslands; fire allowed grasses to invade forests; and the evolution of fire-resistant savanna trees expanded the climate space that savannas can invade., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)