Resource availability and human activity shape the landscape distribution of white rhinoceros, a highly threatened African megaherbivore

The megaherbivore concept suggests that mammals >1000 kg are insensitive to predation as adults. Consequently, their space use should be largely driven by resources. This does not account for the fact that megaherbivores have been hunted by humans for >100,000 years and likely evolved innate responses against human predation. Recent studies indeed show that megaherbivores, such as elephants and rhino, strongly respond to human voices. Few, however, have examined the relative influence of resource versus human risk drivers on the landscape use of megaherbivores. Using a long-term dataset from aerial rhino surveys and poaching events in Hluhluwe-iMfolozi Park, South Africa, we investigated how resource and human risk factors shape the landscape distribution of white rhinoceros (Ceratotherium simum). We used rainfall, fire, catena position, and terrain ruggedness as resource drivers and poaching intensity, and distance to human sound-generating infrastructure as human risk drivers. Both resource and human risk drivers affected rhino landscape distribution. Rhino preferred valley bottoms over midlands and uplands, and the use of the latter two habitats increased during the dry season. During drier wet seasons, rhinos increased their upland habitat use. Rhino avoided the park's fenceline and other infrastructure (roads and camps). Poaching intensity did not influence rhino landscape use. Avoidance of human infrastructures may reduce the effective size of protected areas for rhino. Future work should assess how rhino respond to resources and risk over shorter timescales. Our findings encourage a re-evaluation of the megaherbivore concept to include humans as drivers of their ecology.