Spatial patterns of African ungulate aggregation reveal complex but limited risk effects from reintroduced carnivores

The landscape of fear model, recently advanced in research on the non-lethal effects of carnivores on ungulates, predicts that prey will exhibit detectable antipredator behavior not only during risky times (i.e., predators in close proximity) but also in risky places (i.e., habitat where predators kill prey or tend to occur). Aggregation is an important antipredator response in numerous ungulate species, making it a useful metric to evaluate the strength and scope of the landscape of fear in a multi-carnivore, multi-ungulate system. We conducted ungulate surveys over a 2-year period in South Africa to test the influence of three broad-scale sources of variation in the landscape on spatial patterns in aggregation: (1) habitat structure, (2) where carnivores tended to occur (i.e., population-level utilization distributions), and (3) where carnivores tended to kill ungulate prey (i.e., probabilistic kill site maps). We analyzed spatial variation in aggregation for six ungulate species exposed to predation from recently reintroduced lion (Panthera leo) and spotted hyena (Crocuta crocuta). Although we did detect larger aggregations of ungulates in risky places, these effects existed primarily for smaller-bodied (<150 kg) ungulates and were relatively moderate (change of <= 4 individuals across all habitats). In comparison, ungulate aggregations tended to increase at a slightly lower rate in habitat that was more open. The lion, an ambush (stalking) carnivore, had stronger influence on ungulate aggregation than the hyena, an active (coursing) carnivore. In addition, places where lions tended to kill prey had a greater effect on ungulate aggregation than places where lions tended to occur, but an opposing pattern existed for hyena. Our study reveals heterogeneity in the landscape of fear and suggests broad-scale risk effects following carnivore reintroduction only moderately influence ungulate aggregation size and vary considerably by predator hunting mode, type of predation risk, and prey species.