Feedbacks between soil nutrients and large herbivores in a managed savanna ecosystem

Small-scale fertilization experiments have shown that soil nutrients limit plant productivity in many semiarid grasslands and savannas, but linkages among nutrients, grasses, and grazers are rarely studied in an ecosystem context. We used hectare-scale heterogeneity in soil nutrients created by cattle management practices within a geologically homogeneous savanna to examine relationships among soil nitrogen and phosphorus, aboveground net primary production (ANPP), grass nutrient content, and a mixed community of native and domestic herbivores on central Kenyan rangeland. Increasing soil N and P content was consistently associated with increasing plant productivity and rainfall use efficiency in wet, dry, and drought years. A fertilization experiment and analyses of grass N:P ratios across sites indicated that N is the primary limiting nutrient on nutrient-rich glades, whereas N and P co-limit productivity on nutrient-poor bushland sites. Variation in ANPP among patches within the landscape was linearly correlated with consumption rates of large herbivores. Grazing pressure was consistently high (>60% of ANPP) at all but one site in a dry year (1999), and was greater in nutrient-rich glades (73 +/- 4% of ANPP) than in nutrient-poor bushland sites (43 +/- 7% of ANPP) in a wet year (2001). Grasses of nutrient-rich sites contained sufficient P concentrations to meet requirements for pregnant and lactating ungulates, whereas grasses in nutrient-poor swards were P deficient. Even though native and domestic herbivores selectively used and intensively grazed nutrient-rich sites, productivity on these sites remained high throughout the study. Analyses of nitrogen budgets for nutrient-rich and nutrient-poor sites showed that large herbivores themselves caused a net N input to the former and a net N loss from the latter. Thus, large herbivores not only respond to heterogeneity in soil and plant nutrients across the landscape, but also play a role in maintaining the N-enriched status of highly productive and intensively grazed sites.