Grasses continue to trump trees at soil carbon sequestration following herbivore exclusion in a semiarid African savanna

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 (C-3 vs. C-4) 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 (similar to 22%), woody basal area (similar to 8 m(2)/ha), and woody canopy cover (31%), translating to a similar to 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 similar to 71% of this derived from C-4 grasses (vs. similar to 76% with herbivores present) despite substantial increases in woody cover. We attribute this continued high contribution of C-4 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.