Lions do not change rivers: Complex African savannas preclude top-down forcing by large carnivores

Trophic cascade theories such as the ‘behaviourally-mediated trophic cascade hypothesis’ (BMTCH), have mainstreamed as ecological tools for conserving biodiversity and restoring ecosystems. The BMTCH relies on indirect negative effects of large carnivores through suppression of mesocarnivore activity and habitat use. Importantly, effects of top carnivores on mesocarnivores varies over time and space, is dependent on the species involved, and local context. In South Africa, there are very few free-ranging carnivores, as populations are often restricted to enclosed reserves. While predator-proof fences reduce human-wildlife conflict, they also influence space use within communities. We used an enclosed reserve with a relatively full complement of carnivores to test the generality of the BMTCH in the African context. Using single-species, multi-season occupancy models we investigated the spatial dynamics of multiple carnivores. We also investigated spatial partitioning by vegetation type and temporal partitioning. Our results revealed both support for and against the BMTCH. Lions and spotted hyaenas negatively influenced the detection probability of black-backed jackals and African wildcats, while leopards had a positive effect on these two mesocarnivore species. Additionally, lions positively influenced the detection probability of side-striped jackals. Although space use of carnivores in relation to vegetation type showed minimal evidence of spatial partitioning, each carnivore had a unique combination of abiotic and biotic factors influencing their spatial dynamics, which could facilitate co-existence. Temporal partitioning may also be promoting co-existence as activity patterns of smaller carnivores overlapped the least with sympatric carnivores, particularly lions. Extensive activity overlap between large carnivores does not promote co-existence. We suggest that applying the BMTCH as a universal law across ecosystems is unsupported and may lead to inappropriate conservation and management actions, and prevent protection of ecosystems.