Functional attributes of savanna soils: Contrasting effects of tree canopies and herbivores on bulk density, nutrients and moisture dynamics

Summary: Savannas are highly heterogeneous tree-grass mixtures, and the structural variation imposed by a discontinuous canopy cover results in spatial variation in soil properties such as plant-available nutrients, temperature and soil moisture. Many savannas are also dominated by large vertebrate herbivores, which impose a different suite of effects on soil properties related to consumption, excretion and physical disturbance. In nutrient-poor, water-limited systems, variation in soil resource availability may play a fundamental role in structuring plant communities, but the relative and interactive effects of key drivers of heterogeneity have received little attention in savanna ecosystems. We investigated the independent and interactive effects of herbivores and tree canopy cover on multiple soil properties up to a depth of 10 cm, including bulk density, soil C pools, total and plant-available nutrients, moisture dynamics and temperature at two contrasting sites in Kruger National Park, South Africa. We analysed samples from locations under and between adult tree canopies and inside and outside long-term herbivore exclosures using a nested design. We found strong positive effects of tree canopies on total and labile pools of soil C and N and negative effects on plant-available P and soil temperature. Tree canopy effects on soil moisture availability were small but clearly detectable. During precipitation events, trees increased moisture inputs on granite soils but decreased moisture inputs on basalt soils compared to open sites. Conversely, we found mixed evidence for herbivore effects on soil chemistry and temperature, but a clearer effect on soil moisture, with areas outside exclosures experiencing higher soil moisture retention, particularly immediately following precipitation events. Soil bulk density, a driver of both nutrient and soil moisture dynamics, was lower under trees and when herbivores were excluded. Synthesis. Our results suggest that variation in tree cover is the dominant biotic driver of C, N and P dynamics in these savanna systems in the top 10 cm of soil, that herbivores primarily affect soil moisture content, and that canopy and herbivore effects tend to be additive rather than synergistic.