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Summary Savannas cover a significant fraction of the Earth's land surface. In these ecosystems, C₃trees and C₄grasses coexist persistently, but the mechanisms explaining coexistence remain subject to debate. Different quantitative models have been proposed to explain coexistence, but these models make widely contrasting assumptions about which mechanisms are responsible for savanna persistence. Here, we show that no single existing model fully captures all key elements required to explain tree–grass coexistence across savanna rainfall gradients, but many models make important contributions. We show that recent empirical work allows us to combine many existing elements with new ideas to arrive at a synthesis that combines elements of two dominant frameworks: Walter's two‐layer model and demographic bottlenecks. We propose that functional rooting separation is necessary for coexistence and is the crux of the coexistence problem. It is both well‐supported empirically and necessary for tree persistence, given the comprehensive grass superiority for soil moisture acquisition. We argue that eventual tree dominance through shading is precluded by ecohydrological constraints in dry savannas and by fire and herbivores in wet savannas. Strong asymmetric grass–tree competition for soil moisture limits tree growth, exposing trees to persistent demographic bottlenecks. 

Summary Models of tree–grass coexistence in savannas make different assumptions about the relative performance of trees and grasses under wet vs dry conditions. We quantified transpiration and drought tolerance traits in 26 tree and 19 grass species from the African savanna biome across a gradient of soil water potentials to test for a trade‐off between water use under wet conditions and drought tolerance.We measured whole‐plant hourly transpiration in a growth chamber and quantified drought tolerance using leaf osmotic potential (Ψ_osm). We also quantified whole‐plant water‐use efficiency (WUE) and relative growth rate (RGR) under well‐watered conditions.Grasses transpired twice as much as trees on a leaf‐mass basis across all soil water potentials. Grasses also had a lower Ψ_osmthan trees, indicating higher drought tolerance in the former. Higher grass transpiration and WUE combined to largely explain the threefold RGR advantage in grasses.Our results suggest that grasses outperform trees under a wide range of conditions, and that there is no evidence for a trade‐off in water‐use patterns in wet vs dry soils. This work will help inform mechanistic models of water use in savanna ecosystems, providing much‐needed whole‐plant parameter estimates for African species. 

PremiseBelowground functional traits play a significant role in determining plant water‐use strategies and plant performance, but we lack data on root traits across communities, particularly in the tropical savanna biome, where vegetation dynamics are hypothesized to be strongly driven by tree–grass functional differences in water use. MethodsWe grew seedlings of 21 tree and 18 grass species (N= 5 individuals per species) from the southern African savanna biome under greenhouse conditions and collected fine‐root segments from plants for histological analysis. We identified and measured xylem vessels in 539 individual root cross sections. We then quantified six root vascular anatomy traits and tested them for phylogenetic signals and tree–grass differences in trait values associated with vessel size, number, and hydraulic conductivity. ResultsGrass roots had larger root xylem vessels than trees, a higher proportion of their root cross‐sectional area comprised vessels, and they had higher estimated axial conductivities than trees, while trees had a higher number of vessels per root cross‐sectional area than grasses did. We found evidence of associations between trait values and phylogenetic relatedness in most of these traits across tree species, but not grasses. ConclusionsOur findings support the hypothesis that grass roots have higher water transport capacity than tree roots in terms of maximum axial conductivity, consistent with the observation that grasses are more “aggressive” water users than trees under conditions of high soil moisture availability. Our study identifies root functional traits that may drive differential responses of trees and grasses to soil moisture availability.