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Bastin et al (Reports, 12 May 2017, p. 635) infer forest as more globally extensive than previously estimated using tree cover data. However, their forest definition does not reflect ecosystem function or biotic composition. These structural and climatic definitions inflate forest estimates across the tropics and undermine conservation goals, leading to inappropriate management policies and practices in tropical grassy ecosystems., (Copyright © 2017 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

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-yr 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 semiarid 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., (© 2016 by the Ecological Society of America.)

Contemporary theory on the maintenance and stability of the savanna biome has focused extensively on how climate and disturbances interact to affect tree growth and demography. In particular, the role of fire in reducing tree cover from climatic maxima is now well appreciated, and in certain cases, herbivory also strongly affects tree cover. However, in African savannas and forests, harvest of trees by humans for cooking and heating is an oft overlooked disturbance. Thus, we incorporate tree harvest into a population dynamic model of grasses, savanna saplings, savanna trees, and forest trees. We use assumptions about the differential demographic responses of savanna trees and forest trees to harvest to show how tree harvest influences tree cover, demography, and community composition. Tree harvest can erode the intrinsic basin of attraction for forest and make a state transition via fire to savanna more likely. The savanna state is generally resilient to all but high levels of tree harvest because of the resprouting abilities of savanna trees. In the absence of active fire suppression, our analysis suggests that we can expect to see large and potentially irreversible shifts from forest to savanna as demand increases for charcoal in sub-Saharan Africa. On the other hand, savanna tree species' traits promote savanna stability in the face of low to moderate harvest pressure.

Ecologists have long sought to understand the factors controlling the structure of savanna vegetation. Using data from 2154 sites in savannas across Africa, Australia, and South America, we found that increasing moisture availability drives increases in fire and tree basal area, whereas fire reduces tree basal area. However, among continents, the magnitude of these effects varied substantially, so that a single model cannot adequately represent savanna woody biomass across these regions. Historical and environmental differences drive the regional variation in the functional relationships between woody vegetation, fire, and climate. These same differences will determine the regional responses of vegetation to future climates, with implications for global carbon stocks.

Theoretical models of allometric scaling provide frameworks for understanding and predicting how and why the morphology and function of organisms vary with scale. It remains unclear, however, if the predictions of 'universal' scaling models for vascular plants hold across diverse species in variable environments. Phenomena such as competition and disturbance may drive allometric scaling relationships away from theoretical predictions based on an optimized tree. Here, we use a hierarchical Bayesian approach to calculate tree-specific, species-specific, and 'global' (i.e. interspecific) scaling exponents for several allometric relationships using tree- and branch-level data harvested from three savanna sites across a rainfall gradient in Mali, West Africa. We use these exponents to provide a rigorous test of three plant scaling models (Metabolic Scaling Theory (MST), Geometric Similarity, and Stress Similarity) in savanna systems. For the allometric relationships we evaluated (diameter vs. length, aboveground mass, stem mass, and leaf mass) the empirically calculated exponents broadly overlapped among species from diverse environments, except for the scaling exponents for length, which increased with tree cover and density. When we compare empirical scaling exponents to the theoretical predictions from the three models we find MST predictions are most consistent with our observed allometries. In those situations where observations are inconsistent with MST we find that departure from theory corresponds with expected tradeoffs related to disturbance and competitive interactions. We hypothesize savanna trees have greater length-scaling exponents than predicted by MST due to an evolutionary tradeoff between fire escape and optimization of mechanical stability and internal resource transport. Future research on the drivers of systematic allometric variation could reconcile the differences between observed scaling relationships in variable ecosystems and those predicted by ideal models such as MST.

Savannas are globally important ecosystems of great significance to human economies. In these biomes, which are characterized by the co-dominance of trees and grasses, woody cover is a chief determinant of ecosystem properties. The availability of resources (water, nutrients) and disturbance regimes (fire, herbivory) are thought to be important in regulating woody cover, but perceptions differ on which of these are the primary drivers of savanna structure. Here we show, using data from 854 sites across Africa, that maximum woody cover in savannas receiving a mean annual precipitation (MAP) of less than approximately 650 mm is constrained by, and increases linearly with, MAP. These arid and semi-arid savannas may be considered 'stable' systems in which water constrains woody cover and permits grasses to coexist, while fire, herbivory and soil properties interact to reduce woody cover below the MAP-controlled upper bound. Above a MAP of approximately 650 mm, savannas are 'unstable' systems in which MAP is sufficient for woody canopy closure, and disturbances (fire, herbivory) are required for the coexistence of trees and grass. These results provide insights into the nature of African savannas and suggest that future changes in precipitation may considerably affect their distribution and dynamics.

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Green and senesced leaf nitrogen (N) and phosphorus (P) concentrations of different plant functional groups in savanna communities of Kruger National Park, South Africa were analyzed to determine if nutrient resorption was regulated by plant nutritional status and foliar N:P ratios. The N and P concentrations in green leaves and the N concentrations in senesced leaves differed significantly between the dominant plant functional groups in these savannas: fine-leaved trees, broad-leaved trees and grasses. However, all three functional groups reduced P to comparable and very low levels in senesced leaves, suggesting that P was tightly conserved in this tropical semi-arid savanna ecosystem. Across all functional groups, there was evidence for nutritional control of resorption in this system, with both N and P resorption efficiencies decreasing as green leaf nutrient concentrations increased. However, specific patterns of resorption and the functional relationships between nutrient concentrations in green and senesced leaves varied by nutrient and plant functional group. Functional relationships between N concentrations in green and senesced leaves were indistinguishable between the dominant groups, suggesting that variation in N resorption efficiency was largely the result of inter-life form differences in green leaf N concentrations. In contrast, observed differences in P resorption efficiencies between life forms appear to be the result of both differences in green leaf P concentrations as well as inherent differences between life forms in the fraction of green leaf P resorbed from senescing leaves. Our results indicate that foliar N:P ratios are poor predictors of resorption efficiency in this ecosystem, in contrast to N and P resorption proficiencies, which are more responsive to foliar N:P ratios. [ABSTRACT FROM AUTHOR]

Several explanations for the persistence of tree–grass mixtures in savannas have been advanced thus far. In general, these either concentrate on competition-based mechanisms, where niche separation with respect to limiting resources such as water lead to tree–grass coexistence, or demographic mechanisms, where factors such as fire, herbivory and rainfall variability promote tree–grass persistence through their dissimilar effects on different life-history stages of trees. Tests of these models have been largely site-specific, and although different models find support in empirical data from some savanna sites, enough dissenting evidence exists from others to question their validity as general mechanisms of tree–grass coexistence. This lack of consensus on determinants of savanna structure and function arises because different models: (i) focus on different demographic stages of trees, (ii) focus on different limiting factors of tree establishment, and (iii) emphasize different subsets of the potential interactions between trees and grasses. Furthermore, models differ in terms of the most basic assumptions as to whether trees or grasses are the better competitors. We believe an integration of competition-based and demographic approaches is required if a comprehensive model that explains both coexistence and the relative productivity of the tree and grass components across the diverse savannas of the world is to emerge. As a first step towards this end, we outline a conceptual framework that integrates existing approaches and applies them explicitly to different life-history stage of trees. [ABSTRACT FROM AUTHOR]