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Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open-canopy grassy woodlands) remains limited. To incorporate grasslands into large-scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved - frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human-caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species-diverse plant communities, including endemic species, are slow to recover. Complicating plant-community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re-establish. To guide restoration decisions, we draw on the old-growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old-growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old-growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.

Bastin et al.’s estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration., Funding: Supported by the Texas A&M Sid Kyle Global Savanna Research Initiative (T.W.B.); Swiss National Science Foundation (20FI20_173691) (N.B.); Centre National pour la Recherche Scientifique CNRS PICS 2018-2020 (RESIGRASS) (E.B.); CNPq (Brazil, 303179/2016-3) (G.D.); CNPq (Brazil) (G.W.F.); CNPq (Brazil, 303988/2018-5) (A.F.); NASA award NNX17AK14G (F.F.); NSF award 1354943 (W.A.H.); Fundação de Amparo à Pesquisa do Estado de Minas Gerais (Brazil, 2016/13232-5) (S.L.S.); the Office of the Royal Society (IC170015) (C.E.R.L.); CNPq (Brazil, 310345/2018-9) (G.E.O.); the Spanish Government (FIROTIC, PGC2018-096569-B-I00) (J.G.P.); the National Research Foundation (ACCESS, 114695) (N.S.); CNPq (Brazil, 303568/2017-8) (F.A.O.S.); NSF awards 1342703 and 1926431 (C.J.S. and D.M.G.); NSF award EAR-1253713 (C.A.E.S.); Deutsche Forschungsgemeinschaft grant 5579 POEM (V.M.T.); and USDA-NIFA Sustainable Agricultural Systems Grant 12726253 (J.W.V.)

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open-canopy grassy woodlands) remains limited. To incorporate grasslands into large-scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved - frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human-caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species-diverse plant communities, including endemic species, are slow to recover. Complicating plant-community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re-establish. To guide restoration decisions, we draw on the old-growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old-growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old-growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.

Deforestation as a result of burning and land conversion in the tropics and subtropics has been widely studied and active restoration of forests has been widely promoted. Besides other benefits, reforestation can sequester carbon thereby reducing CO 2 emissions to the atmosphere. However, before grasslands are targeted for ‘reforestation', it is necessary to distinguish whether they are ancient natural grasslands or secondary vegetation colonizing deforested areas. Here we report the results of a study comparing primary grasslands in South Africa with 4–40 year old secondary grasslands recovering from afforestation with Pinus species. Primary grasslands had significantly higher plant species richness overall, especially of forb species. Ground cover of primary grasslands was more evenly distributed among species than secondary grasslands which tended to mono-dominance. Forbs with underground storage organs (USOs) were common in primary grasslands but conspicuously absent in the recovering systems. Comparison of secondary grasslands of different ages (up to 40 years) showed negligible recovery of the original species composition. Three key features distinguish old growth primary from secondary grasslands: total and forb species numbers, evenness of species contributions to cover and the presence of USOs. Old growth grasslands also differed in their fire response, showing significant post-burn resprouting and fire-stimulated flowering in contrast to secondary grasslands. Though similar contrasting attributes of ancient and secondary grasslands have been reported in South America, more studies are needed to explore their generality in other geographical regions. This article is part of the themed issue ‘Tropical grassy biomes: linking ecology, human use and conservation’.

Africa has the most extensive C4 grassy biomes of any continent. They are highly flammable accounting for greater than 70% of the world's burnt area. Much of Africa's savannas and grasslands occur in climates warm enough and wet enough to support closed forests. The combination of open grassy systems and the frequent fires they support have long been interpreted as anthropogenic artefacts caused by humans igniting frequent fires. True grasslands, it was believed, would be restricted to climates too dry or too cold to support closed woody vegetation. The idea that higher-rainfall savannas are anthropogenic and that fires are of human origin has led to initiatives to ‘reforest’ Africa's open grassy systems paid for by carbon credits under the assumption that the net effect of converting these system to forests would sequester carbon, reduce greenhouse gases and mitigate global warming. This paper reviews evidence for the antiquity of African grassy ecosystems and for the fires that they sustain. Africa's grassy biomes and the fires that maintain them are ancient and there is no support for the idea that humans caused large-scale deforestation. Indicators of old-growth grasslands are described. These can help distinguish secondary grasslands suitable for reforestation from ancient grasslands that should not be afforested. This article is part of the themed issue ‘The interaction of fire and mankind’.

The coastal grasslands in north-eastern South Africa are a severely threatened vegetation type rich in plant species, particularly forbs. Many of the forbs have underground storage organs which allow them to resprout rapidly after fires. A significant portion of this land was placed under commercial pine afforestation in the 1950s. The pine plantations have since been removed starting 17 years ago and restored to grasslands within a conservation area. We assessed the effects of plantations on grassland plant diversity and functional trait composition by sampling 64 circular quadrats of 5 m radius distributed equally in restored versus natural grasslands. The difference in plant diversity was dramatic with the natural grassland supporting 221 species of which 163 were forbs compared with 144 and only 73 forb species in restored grasslands. Major differences in species composition were recorded, especially for forb species. Natural grasslands were dominated by resprouters (130 species) but these were rare in the restored grasslands (36 species). Differences in plant species response to fire were also evident for the two grassland states. In contrast to coastal forest restoration studies in the same area which have shown near linear increases in woody species with time, restored grasslands showed no increase in forb species richness with increasing time since clear-felling of pines. Our results indicate that current methods for restoring these grasslands are inadequate and that restoring grasslands may be a lot harder than previously thought. Considerable effort should be made in conserving what is left of natural grasslands.

International audience; We expand the concept of “old growth” to encompass the distinct ecologies and conservation values of the world's ancient grass-dominated biomes. Biologically rich grasslands, savannas, and open-canopy woodlands suffer from an image problem among scientists, policy makers, land managers, and the general public, that fosters alarming rates of ecosystem destruction and degradation. These biomes have for too long been misrepresented as the result of deforestation followed by arrested succession. We now know that grassy biomes originated millions of years ago, long before humans began deforesting. We present a consensus view from diverse geographic regions on the ecological characteristics needed to identify old-growth grasslands and to distinguish them from recently formed anthropogenic vegetation. If widely adopted, the old-growth grassland concept has the potential to improve scientific understanding, conservation policies, and ecosystem management.