Topo-edaphic controls over woody plant biomass in South African savannas

The distribution of woody biomass in savannas reflects spatial patterns fundamental to ecosystem processes, such as water flow, competition, and herbivory, and is a key contributor to savanna ecosystem services, such as fuelwood supply. While total precipitation sets an upper bound on savanna woody biomass, the extent to which substrate and terrain constrain trees and shrubs below this maximum remains poorly understood, often occluded by local-scale disturbances such as fire and trampling. Here we investigate the role of hillslope topography and soil properties in controlling woody plant aboveground biomass (AGB) in Kruger National Park, South Africa. Large-area sampling with airborne Light Detection and Ranging (LiDAR) provided a means to average across local-scale disturbances, revealing an unexpectedly linear relationship between AGB and hillslopeposition on basalts, where biomass levels were lowest on crests, and linearly increased toward streams (R-2 = 0.91). The observed pattern was different on granite substrates, where AGB exhibited a strongly non-linear relationship with hillslope position: AGB was high on crests, decreased midslope, and then increased near stream channels (R-2 = 0.87). Overall, we observed 5-to-8-fold lower AGB on clayey, basalt-derived soil than on granites, and we suggest this is due to herbivore-fire interactions rather than lower hydraulic conductivity or clay shrinkage/swelling, as previously hypothesized. By mapping AGB within and outside fire and herbivore exclosures, we found that basalt-derived soils support tenfold higher AGB in the absence of fire and herbivory, suggesting high clay content alone is not a proximal limitation on AGB. Understanding how fire and herbivory contribute to AGB heterogeneity is critical to predicting future savanna carbon storage under a changing climate.