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1. The stress-gradient hypothesis (SGH) predicts an increasing importance of facilitative mechanisms relative to competition along gradients of increasing environmental stress. Although developed across a variety of ecosystems, the SGH's relevance to the dynamic tree—grass systems of global savannas remains unclear. Here, we present a meta-analysis of empirical studies to explore emergent patterns of tree—grass relationships in global savannas in the context of the SGH. 2. We quantified the net effect of trees on understorey grass production relative to production away from tree canopies along a rainfall gradient in tropical and temperate savannas and compared these findings to the predictions of the SGH. We also analysed soil and plant nutrient concentrations in subcanopy and open-grassland areas to investigate the potential role of nutrients in determining grass production in the presence and absence of trees. 3. Our meta-analysis revealed a shift from net competitive to net facilitative effects of trees on subcanopy grass production with decreasing annual precipitation, consistent with the SGH. We also found a significant difference between sites from Africa and North America, suggesting differences in tree—grass interactions in the savannas of tropical and temperate regions. 4. Nutrient analyses indicate no change in nutrient ratios along the rainfall gradient, but consistent nutrient enrichment under tree canopies. 5. Synthesis. Our results help to resolve questions about the SGH in semi-arid systems, demonstrating that in mixed tree—grass systems, trees facilitate grass growth in drier regions and suppress grass growth in wetter regions. Relationships differ, however, between African and North American sites representing tropical and temperate bioclimates, respectively. The results of this meta-analysis advance our understanding of tree—grass interactions in savannas and contribute a valuable data set to the developing theory behind the SGH.

Management of tree cover, either to curb bush encroachment or to mitigate losses of woody cover to over-browsing, is a major concern in savanna ecosystems. Once established, trees are often “trapped” as saplings, since interactions among disturbance, plant competition, and precipitation delay sapling recruitment into adult size classes. Saplings can be directly suppressed by wildlife browsing and competition from adjacent plants, and indirectly facilitated by grazers, such as cattle, which feed on neighboring grasses. Yet few experimental studies have simultaneously quantified the effects of cattle and wildlife on sapling growth, particularly over long time scales. We used a series of replicated 4-ha herbivore-manipulation plots to investigate the net effects of wildlife and moderate cattle grazing on Acacia drepanolobium sapling growth over 10 years that encompassed extended wet and dry periods. We also simulated more intense cattle grazing using grass removal treatments (0.5-m radius around saplings), and we quantified the role of intraspecific tree competition using neighborhood tree surveys (trees within a 3-m radius). Wildlife, which included elephants, had a positive effect on sapling growth. Wildlife also reduced neighbor tree density during the 10-yr study, which likely caused the positive effect of wildlife on saplings. Although moderate cattle grazing did not affect sapling growth, grass removal treatments simulating heavy grazing increased sapling growth. Both grass removal and neighbor tree effects on saplings were strongest during above-average rainfall years following drought. This highlights that livestock-driven reductions in grass cover and catastrophic wildlife damage to trees during droughts present a need, or an opportunity, for targeted management of sapling growth and woody plant cover during ensuing wet periods.

During the past century, the biomass of woody species has increased in many grassland and savanna ecosystems. As many of these species fix nitrogen symbiotically, they may alter not only soil nitrogen (N) conditions but also those of phosphorus (P). We studied the N‐fixing shrub Dichrostachys cinerea in a mesic savanna in Zambia, quantifying its effects upon pools of soil N, P, and carbon (C), and availabilities of N and P. We also evaluated whether these effects induced feedbacks upon the growth of understory vegetation and encroaching shrubs. Dichrostachys cinerea shrubs increased total N and P pools, as well as resin‐adsorbed N and soil extractable P in the top 10‐cm soil. Shrubs and understory grasses differed in their foliar N and P concentrations along gradients of increasing encroachment, suggesting that they obtained these nutrients in different ways. Thus, grasses probably obtained them mainly from the surface upper soil layers, whereas the shrubs may acquire N through symbiotic fixation and probably obtain some of their P from deeper soil layers. The storage of soil C increased significantly under D. cinerea and was apparently not limited by shortages of either N or P. We conclude that the shrub D. cinerea does not create a negative feedback loop by inducing P‐limiting conditions, probably because it can obtain P from deeper soil layers. Furthermore, C sequestration is not limited by a shortage of N, so that mesic savanna encroached by this species could represent a C sink for several decades. We studied the effects of woody encroachment on soil N, P, and C pools, and availabilities of N and P to Dichrostachys cinerea shrubs and to the understory vegetation. Both N and P pools in the soil increased along gradients of shrub age and cover, suggesting that N fixation by D. cinerea did not reduce the P supply. This in turn suggests that continued growth and carbon sequestration in this mesic savanna ecosystems are unlikely to be constrained by nutrient limitation and could represent a C sink for several decades.

Savanna ecosystems, defined by the codominance of trees and grasses, cover one-fifth of the world's land surface and are of great socioeconomic and biological importance. Yet, the fundamental question of how trees and grasses coexist to maintain the savanna state remains poorly understood. Many models have been put forward to explain tree—grass coexistence, but nearly all have assumed that grasses do not limit tree growth and demography beyond the sapling stage. This assumption, however, has rarely been tested. Here I show that grass can strongly suppress the growth of trees. I removed grass around trees of three size classes in an Acacia drepanolobium savanna in Laikipia, Kenya. For even the largest trees, grass removal led to a doubling in growth and a doubling in the probability of transitioning to the next size class over two years. These results suggest that grass competition in productive (nutrient-rich) savannas may limit tree growth as much as herbivory and fire (the main factors thought to determine tree demography within a rainfall region) and should be incorporated into savanna models if tree—grass coexistence and savanna dynamics are to be understood.

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant “bottom-up” effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating “top-down” effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the “landscape of fear” in which African savanna herbivores exist.

1. A recent meta-analysis suggested that differences in rainfall are a cause of variation in tree-grass interactions in savannas, with trees facilitating growth of understorey grasses in low-rainfall areas, but competing with them under higher rainfall. We hypothesized that this effect of rainfall upon understorey productivity is modified by differences in the growth form of the woody plants (i.e. the height of the lower canopy) or by their capacity to fix nitrogen. 2. We performed a meta-analysis of the effects of woody plants on understorey productivity, incorporating canopy height and N-fixation, and their interaction with rainfall. 3. N-fixing woody plants enhanced understorey productivity, whereas non-fixers had a neutral or negative effect, depending on high or low canopy, respectively. We found a strong negative correlation between rainfall and the degree to which trees enhanced understorey productivity, but only for trees with a high canopy. 4. Synthesis. The effect of woody plants on understorey productivity depends not only on rainfall, but also on their growth form and their capacity to fix N. Facilitation occurs mostly when woody plants ameliorate both water and nitrogen conditions. However, a low canopy suppresses understorey vegetation by competing for light, regardless of water and nutrient relations.

Vertebrate herbivores and fire are known to be important drivers of vegetation dynamics in African savannas. It is of particular importance to understand how changes in herbivore population density, especially of elephants, and fire frequency will affect the amount of tree cover in savanna ecosystems, given the critical importance of tree cover for biodiversity, ecosystem function, and human welfare. We developed a spatially realistic simulation model of vegetation, fire, and dominant herbivore dynamics, tailored to the Serengeti ecosystem of east Africa. The model includes key processes such as tree—grass competition, fire, and resource-based density dependence and adaptive movement by herbivores. We used the model to project the ecosystem 100 years into the future from its present state under different fire, browsing (determined by elephant population density), and grazing (with and without wildebeest present) regimes. The model produced the following key results: (1) elephants and fire exert synergistic negative effects on woody cover; when grazers are excluded, the impact of fire and the strength of the elephant—fire interaction increase; (2) at present population densities of 0.15 elephants/km2, the total amount of woody cover is predicted to remain stable in the absence of fire, but the mature tree population is predicted to decline regardless of the fire regime; without grazers present to mitigate the effects of fire, the size structure of the tree population will become dominated by seedlings and mature trees; (3) spatial heterogeneity in tree cover varies unimodally with elephant population density; fire increases heterogeneity in the presence of grazers and decreases it in their absence; (4) the marked rainfall gradient in the Serengeti directly affects the pattern of tree cover in the absence of fire; with fire, the woody cover is determined by the grazing patterns of the migratory wildebeest, which are partly rainfall driven. Our results show that, in open migratory ecosystems such as the Serengeti, grazers can modulate the impact of fire and the strength of the interaction between fire and browsers by altering fuel loads and responding to the distribution of grass across the landscape, and thus exert strong effects on spatial patterns of tree cover.

We present a continental-scale analysis that explores the processes controlling woody community structure in tropical savannas. We analyse how biotic and abiotic factors interact to promote and modify tree cover, examine alternative ecological hypotheses and quantify disturbance effects using satellite estimates of tree cover. African savannas. Tree cover is represented as a resource-driven potential cover related to rainfall and soil characteristics perturbed by natural and human factors such as fire, cattle grazing, human population and cultivation. Within this framework our approach combines semi-empirical modelling and information theory to identify the best models. Woody community structure across African savannas is best represented by a sigmoidal response of tree cover to mean annual precipitation (MAP), with a dependency on soil texture, which is modified by the separate effects of fire, domestic livestock, human population density and cultivation intensity. This model explains c. 66% of the variance in tree cover and appears consistent across the savanna regions of Africa. The analysis provides a new understanding of the importance and interaction of environmental and disturbance factors that create the broad spatial patterns of tree cover observed in African savannas. Woody cover increases with rainfall, but is modified by disturbances. These 'perturbation' effects depend on MAP regimes: in arid savannas (MAP < 400 mm) they are generally small (< 1% decrease in cover), while in semi-arid and mesic savannas (400-1600 mm), perturbations result in an average 2% (400 mm) to 23% (1600 mm) decrease in cover; fire frequency and human population have more influence than cattle, and cultivation appears, on average, to lead to small increases in woody cover. Wet savannas (1600-2200 mm) are controlled by perturbations that inhibit canopy closure and reduce tree cover by, on average, 24-34%. Full understanding of the processes determining savanna structure requires consideration of resource limitation and disturbance dynamics.

1. Savanna ecosystems - defined by the coexistence of trees and grasses - cover more than one-fifth the world's land surface and harbour most of the world's rangelands, livestock and large mammal diversity. Savanna trees can have a variety of effects on grasses, with consequences for the wild and domestic herbivores that depend on them. 2. Studies of these effects have focused on two different spatial scales. At the scale of individual trees, many studies have shown net positive effects of trees on sub-canopy grass nutrient concentrations and biomass. At the landscape scale, other studies have shown negative effects of high tree densities on grass productivity. These disparate results have led to different conclusions about the effects of trees on forage quality and ungulate nutrition in savannas. 3. We integrate these approaches by examining the effects of trees on grasses at both spatial scales and across a range of landscape-scale tree densities. We quantified grass biomass, species composition and nutrient concentrations in these different contexts in an Acacia drepanolobium savanna in Laikipia, Kenya. 4. Individual trees had positive effects on grass biomass, most likely because trees enrich soil nitrogen. Grass leaf phosphorus in sub-canopy areas, however, was depressed. The effects of individual trees could explain the effects of increasing landscape-scale tree cover for the biomass of only two of the four dominant grass species. 5. The negative effects of trees on grass and soil phosphorus, combined with depressed grass productivity in areas of high tree cover, suggest that ungulate nutrition may be compromised in areas with many trees. 6. Synthesis. We conclude that few, isolated trees may have positive local effects on savanna grasses and forage, but in areas of high tree density the negative landscape-scale effects of trees are likely to outweigh these positive effects. In savannas and other patchy landscapes, attempts to predict the consequences of changes in patch abundances for ecosystem services (e.g. rangeland productivity and carbon sequestration) will depend on our understanding of the extent to which local, patch-scale dynamics do or do not predict landscape-scale dynamics.

Nutrient enrichment is increasingly affecting many tropical ecosystems, but there is no information on how this affects tree biodiversity. To examine dynamics in vegetation structure and tree species biomass and diversity, we annually remeasured tree species before and for six years after repeated additions of nitrogen (N) and phosphorus (P) in permanent plots of abandoned pasture in Amazonia. Nitrogen and, to a lesser extent, phosphorus addition shifted growth among woody species. Nitrogen stimulated growth of two common pioneer tree species and one common tree species adaptable to both high- and low-light environments, while P stimulated growth only of the dominant pioneer tree Rollinia exsucca (Annonaceae). Overall, N or P addition reduced tree assemblage evenness and delayed tree species accrual over time, likely due to competitive monopolization of other resources by the few tree species responding to nutrient enrichment with enhanced establishment and/or growth rates. Absolute tree growth rates were elevated for two years after nutrient addition. However, nutrient-induced shifts in relative tree species growth and reduced assemblage evenness persisted for more than three years after nutrient addition, favoring two nutrient-responsive pioneers and one early-secondary tree species. Surprisingly, N + P effects on tree biomass and species diversity were consistently weaker than N-only and P-only effects, because grass biomass increased dramatically in response to N + P addition. The resulting intensified competition probably prevented an expected positive N + P synergy in the tree assemblage. Thus, N or P enrichment may favor unknown tree functional response types, reduce the diversity of coexisting species, and delay species accrual during structurally and functionally complex tropical rainforest secondary succession.