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Although studies have shown that mammalian herbivores often limit aboveground carbon storage in savannas, their effects on belowground soil carbon storage remain unclear. Using three sets of long‐term, large herbivore exclosures with paired controls, we asked how almost two decades of herbivore removal from a semiarid savanna in Laikipia, Kenya affected aboveground (woody and grass) and belowground soil carbon sequestration, and determined the major source (C3 vs. C4) of belowground carbon sequestered in soils with and without herbivores present. Large herbivore exclusion, which included a diverse community of grazers, browsers, and mixed‐feeding ungulates, resulted in significant increases in grass cover (~22%), woody basal area (~8 m2/ha), and woody canopy cover (31%), translating to a ~8.5 t/ha increase in aboveground carbon over two decades. Herbivore exclusion also led to a 54% increase (20.5 t/ha) in total soil carbon to 30‐cm depth, with ~71% of this derived from C4 grasses (vs. ~76% with herbivores present) despite substantial increases in woody cover. We attribute this continued high contribution of C4 grasses to soil C sequestration to the reduced offtake of grass biomass with herbivore exclusion together with the facilitative influence of open sparse woody canopies (e.g., Acacia spp.) on grass cover and productivity in this semiarid system.

1. Forest–grassland mosaics, with abrupt boundaries between the two vegetation types, occur across the globe. Fire and herbivory are widely considered primary drivers that maintain these mosaics by limiting tree establishment in grasslands, while edaphic factors and frosts are generally considered to be secondary factors that reinforce these effects. However, the relative importance of these drivers likely varies across systems. In particular, although frost is known to occur in many montane tropical mosaics, experimental evidence for its role as a driving factor is limited. 2. We used replicated in situ transplant and warming experiments to examine the role of microclimate (frost and freezing temperatures) and soil in influencing germination and seedling survival of both native forest trees and alien invasive Acacia trees in grasslands of a tropical montane forest–grassland mosaic in the Western Ghats of southern India. 3. Seed germination of both native and alien tree species was higher in grasslands regardless of soil type, indicating that germination was not the limiting stage to tree establishment. However, irrespective of soil type, native seedlings in grasslands incurred high mortality following winter frosts and freezing temperatures relative to native seedlings in adjoining forests where freezing temperatures did not occur. Seedling survival through the tropical winter was thus a primary limitation to native tree establishment in grasslands. In contrast, alien Acacia seedlings in grasslands incurred much lower levels of winter mortality. Experimental night‐time warming in grasslands significantly enhanced over‐winter survival of all tree seedlings, but increases were much greater for alien Acacia than for native tree seedlings. 4. Synthesis. Our results provide evidence for a primary role for frost and freezing temperatures in limiting tree establishment in grasslands of this tropical montane forest–grassland mosaic. Future increases in temperature are likely to release trees from this limitation and favour tree expansion into grasslands, with rates of expansion of non‐native Acacia likely to be much greater than that of native trees. We suggest that studies of frost limitation to plant establishment are needed across a range of tropical ecosystems to re‐evaluate the general importance of frost as a driver of vegetation transitions in the tropics.

Bark damage resulting from elephant feeding is common in African savanna trees with subsequent interactions with fire, insects, and other pathogens often resulting in tree mortality. Yet, surprisingly little is known about how savanna trees respond to bark damage. We addressed this by investigating how the inner bark of marula (Sclerocarya birrea), a widespread tree species favoured by elephants, recovers after bark damage. We used a long‐term fire experiment in the Kruger National Park to measure bark recovery with and without fire. At 24 months post‐damage, mean wound closure was 98, 92, and 72%, respectively, in annual and biennial burns and fire‐exclusion treatments. Fire exclusion resulted in higher rates of ant colonization of bark wounds, and such ant colonization resulted in significantly lower bark recovery. We also investigated how ten common savanna tree species respond to bark damage and tested for relationships between bark damage, bark recovery, and bark traits while accounting for phylogeny. We found phylogenetic signal in bark dry matter content, bark N and bark P, but not in bark thickness. Bark recovery and damage was highest in species which had thick moist inner bark and low wood densities (Anacardiaceae), intermediate in species which had moderate inner bark thickness and wood densities (Fabaceae) and lowest in species which had thin inner bark and high wood densities (Combretaceae). Elephants prefer species with thick, moist inner bark, traits that also appear to result in faster recovery rates.

Recent work suggests that savanna woody plant species utilise two different strategies based on their defences against herbivory; a low nutrient/high chemical defence strategy and a nutrition paired with mostly architectural defences strategy. The concept that chemical and structural defences can augment each other and do not necessarily trade-off has emanated from this work. In this study, we examine woody plant defence strategies, how these respond to herbivore removal and how they affect plant growth in an East African savanna. At three paired long-term exclosure sites with high browser and mixed-feeder densities at Mpala Ranch, Kenya, we investigated: (a) whether defences employed by the dominant fine- and broad-leaved woody savanna species form defence strategies and if these align with previously proposed strategies, (b) how nine key plant defence traits respond to herbivore removal and (c) how effective the different defence strategies are at protecting against intense herbivory (by measuring plant growth with and without herbivores present). We identified three defence strategies. We found a group (a) with high N, short spines and high N-free secondary metabolites, a group (b) with high N, long spines and low N-free secondary metabolites and a group (c) with moderate N, no spines and low N-free secondary metabolites (most likely defended by unmeasured chemical defences). Structural defences (spine length, branching) were generally found to be induced by herbivory, leaf available N increased or did not respond, and N-free secondary metabolites decreased or did not respond to herbivory. Species with long spines combined with increased “caginess” (dense canopy architecture arising from complex arrangement of numerous woody and spiny axis categories) of branches, maintained the highest growth under intense browsing, compared to species with short spines and high N-free secondary metabolites and species with no spines and low N-free secondary metabolites. Synthesis. At our study site, structural traits (i.e. spines, increased caginess) were the most inducible and effective defences against intense mammalian herbivory. We propose that high levels of variability in the way that nutrient and defence traits combine may contribute to the coexistence of closely related species comprising savanna woody communities.

Two dominant biomes that occur across the southern Indian peninsula are dry deciduous “forests” and evergreen forests, with the former occurring in drier regions and the latter in wetter regions, sometimes in close proximity to each other. Here we compare stem and leaf traits of trees from multiple sites across these biomes to show that dry deciduous “forest” species have, on average, lower height: diameter ratios, lower specific leaf areas, higher wood densities and higher relative bark thickness, than evergreen forest species. These traits are diagnostic of these dry deciduous “forests” as open, well-lit, drought-, and fire-prone habitats where trees are conservative in their growth strategies and invest heavily in protective bark tissue. These tree traits together with the occurrence of a C4 grass-dominated understory, diverse mammalian grazers, and frequent fires indicate that large tracts of dry deciduous “forests” of southern India are more accurately classified as mesic deciduous “savannas.”

The success of invasive species eradication depends on a variety of factors, including those that initially facilitated the invasion, as well as removal and post-removal protocols. Two factors that appear to influence invasion by, and eradication of, the Neotropical shrub Lantana camara (L.), in southern Indian deciduous forests, are rainfall and removal method. However, their role in influencing eradication success is yet to be quantified, and remains unclear. We conducted an experiment to clarify how rainfall (high vs. low) and removal method (cutting vs. uprooting Lantana) influence re-invasion by Lantana, and native plant recovery. Rainfall influenced both eradication effort and outcomes—drier forest had lower starting levels of invader biomass, requiring less initial eradication effort, as well as lower subsequent Lantana re-invasion (from seed and rootstock) whereas wetter forest typically had greater starting levels of invader biomass, requiring considerably greater initial eradication effort, and greater Lantana re-invasion. However, wetter forest also showed greater native tree and forb recovery. Therefore, the availability of funds, local environmental gradients, and restoration priorities should inform the selection of restoration sites. With regard to removal method, uprooting combined with weeding of germinating Lantana, particularly after the rainy season, minimized overall re-invasion. Therefore, uprooting, followed by regular weeding of germinating Lantana and secondary invaders, is crucial to long-term Lantana eradication success.

Colonial encounters with tropical ecosystems were primarily driven by profit-oriented management practices; witness the extensive network of timber and forestry practices that were set up across colonial India. In contrast, the colonial engagement with the montane forest-grassland mosaics of the higher reaches of the Western Ghats in southern India was marked by intensive investment in vegetation management by colonial foresters that yielded no profits. In this archival study, we trace the history of extensive vegetation transformation in this landscape from the early nineteenth to the early twentieth century. We show how the misperception that the grasslands within this mosaic must have resulted from tree felling, fire-setting and buffalo grazing by indigenous communities led colonial foresters into a century-long effort at ‘foresting’ the grasslands, primarily through large-scale planting of exotic tree species. These efforts persisted despite economic losses and ecological evidence that native tree seedlings planted in the grasslands repeatedly failed to establish. These policies continued unabated into the late twentieth century in newly independent India. Today, the once picturesque landscapes of these ancient forest-grassland mosaics are diminished by large-scale plantations of exotic species. Some of these species have become invasive and pose significant threats to the remnant natural grasslands. While this historical narrative is set in the forest-grassland mosaics of southern India, it finds striking parallels in the current day, with grasslands and savannas globally threatened by the misperception that they are ‘degraded ecosystems’ that can be ‘forested’ or converted to other ‘productive’ land uses. We suggest that this case history portends the potential fates of many of earth's threatened tropical grasslands and savannas.

Tropical semi-arid grasslands are a widespread and ecologically and economically important terrestrial biome. Here, we use paleoecology to understand woodland–grassland transitions across the mid- to late-Holocene period in the Banni grassland, western India. Multi proxy analyses involving palynology, phytoliths and elemental geochemistry were carried out on two sediment cores retrieved from wetlands (Chachi and Luna), to understand temporal fluctuations in vegetation, moisture availability and other environmental parameters. Based on the results, the Chachi core was divided into two major climatic phases. Phase 1 (4600–2500 cal. yr BP) was characterised by high precipitation and abundance of pollen types and phytolith morphotypes that indicate the presence of woody savanna, and mesic herbaceous taxa. Phase 2 (2500 cal. yr BP to the present) was characterised by lower precipitation, lower abundance of mesic taxa and an increase in grass phytolith abundance. However, the period from ~1000 cal. yr BP to the present was characterised by the increased abundance of leguminous taxa, dryland herbs/shrubs and a decline in grass phytolith abundance. The Luna core (~1000 cal. yr BP to the present) also showed results matching with the Chachi core for this latter period. Overall, moisture availability in the ecosystem appears to have declined since 4600 cal. yr BP, and the vegetation has responded to this. Although the balance between tree, shrub and grass elements has fluctuated, overall, the region has remained as an open ‘grass and shrub savanna’ with sparse woody vegetation throughout this period. Our study provides insights into the vegetation dynamics and environmental settings in a poorly understood tropical arid-grassland ecosystem from Asia during the mid-late-Holocene.

African savannas support an iconic fauna, but they are undergoing large-scale population declines and extinctions of large (>5 kg) mammals. Long-term, controlled, replicated experiments that explore the consequences of this defaunation (and its replacement with livestock) are rare. The Mpala Research Centre in Laikipia County, Kenya, hosts three such experiments, spanning two adjacent ecosystems and environmental gradients within them: the Kenya Long-Term Exclosure Experiment (KLEE; since 1995), the Glade Legacies and Defaunation Experiment (GLADE; since 1999), and the Ungulate Herbivory Under Rainfall Uncertainty experiment (UHURU; since 2008). Common themes unifying these experiments are (1) evidence of profound effects of large mammalian herbivores on herbaceous and woody plant communities; (2) competition and compensation across herbivore guilds, including rodents; and (3) trophic cascades and other indirect effects. We synthesize findings from the past two decades to highlight generalities and idiosyncrasies among these experiments, and highlight six lessons that we believe are pertinent for conservation. The removal of large mammalian herbivores has dramatic effects on the ecology of these ecosystems; their ability to rebound from these changes (after possible refaunation) remains unexplored.

Tropical grasslands and savannas are globally extensive, and are of significant environmental, economic, and ecological importance. These ecosystems are anticipated to be particularly sensitive to future changes in climate, and understanding how these systems have responded to climatic changes in the past can provide us with insights into their potential responses to future global change. In this study, the temporal dynamics of C 3 -C 4 vegetation changes in response to changes in moisture availability, local fire events and changing levels of herbivory in a summer-rainfall region of Western India are reconstructed for the past ~4600calyrBP. Paleodata such as stable carbon isotope of bulk organic matter (δ 13 C org ), oxygen isotope from carbonate shells (δ 18 O shell ), macro-charcoal and herbivore dung fungal spores are reported from the retrieved cores of two wetland sites located in the Banni grasslands of Western India. Results show that vegetation in the Banni was composed mostly of C 3 vegetation from ~4600 to ~2500calyrBP, after which there was a decline in C 3 vegetation. From the late-Holocene to the present, there was a mix of both C 3 and C 4 vegetation, with C 4 grasses being more abundant in the ecosystem. These shifts were coincident with rainfall changes from more mesic conditions during ~4600 to ~2500calyrBP to more arid conditions towards the present as indicated by δ 18 O shell isotope data. The period of increase in C 4 vegetation also coincides with a period of increased biotic disturbances in the ecosystem, particularly fire. Given the current scenarios of global warming, recurrent drought events and increased anthropogenic use of similar ecosystems, such studies can provide us insights into potential future trajectories of these ecosystems.

The majority of research on savanna vegetation dynamics has focused on the coexistence of woody and herbaceous vegetation. Interactions among woody plants in savannas are relatively poorly understood. We present data from a 10-year longitudinal study of spatially explicit growth patterns of woody vegetation in an East African savanna following exclusion of large herbivores and in the absence of fire. We examined plant spatial patterns and quantified the degree of competition among woody individuals. Woody plants in this semi-arid savanna exhibit strongly clumped spatial distributions at scales of 1 - 5 m. However, analysis of woody plant growth rates relative to their conspecific and heterospecific neighbors revealed evidence for strong competitive interactions at neighborhood scales of up to 5 m for most woody plant species. Thus, woody plants were aggregated in clumps despite significantly decreased growth rates in close proximity to neighbors, indicating that the spatial distribution of woody plants in this region depends on dispersal and establishment processes rather than on competitive, density-dependent mortality. However, our documentation of suppressive effects of woody plants on neighbors also suggests a potentially important role for tree-tree competition in controlling vegetation structure and indicates that the balanced-competition hypothesis may contribute to well-known patterns in maximum tree cover across rainfall gradients in Africa.