title: In the absence of a "landscape of fear": How lions, hyenas, and cheetahs coexist.
abstract: Aggression by top predators can create a "landscape of fear" in which subordinate predators restrict their activity to low-risk areas or times of day. At large spatial or temporal scales, this can result in the costly loss of access to resources. However, fine-scale reactive avoidance may minimize the risk of aggressive encounters for subordinate predators while maintaining access to resources, thereby providing a mechanism for coexistence. We investigated fine-scale spatiotemporal avoidance in a guild of African predators characterized by intense interference competition. Vulnerable to food stealing and direct killing, cheetahs are expected to avoid both larger predators; hyenas are expected to avoid lions. We deployed a grid of 225 camera traps across 1,125 km2in Serengeti National Park, Tanzania, to evaluate concurrent patterns of habitat use by lions, hyenas, cheetahs, and their primary prey. We used hurdle models to evaluate whether smaller species avoided areas preferred by larger species, and we used time-to-event models to evaluate fine-scale temporal avoidance in the hours immediately surrounding top predator activity. We found no evidence of long-term displacement of subordinate species, even at fine spatial scales. Instead, hyenas and cheetahs were positively associated with lions except in areas with exceptionally high lion use. Hyenas and lions appeared to actively track each, while cheetahs appear to maintain long-term access to sites with high lion use by actively avoiding those areas just in the hours immediately following lion activity. Our results suggest that cheetahs are able to use patches of preferred habitat by avoiding lions on a moment-to-moment basis. Such fine-scale temporal avoidance is likely to be less costly than long-term avoidance of preferred areas: This may help explain why cheetahs are able to coexist with lions despite high rates of lion-inflicted mortality, and highlights reactive avoidance as a general mechanism for predator coexistence.
authors.affiliation: Department of Ecology Evolution and Behavior University of Minnesota Saint Paul MN USA; Department of Physics University of Oxford Oxford UK.
authors.forename: Alexandra
authors.initials: A
authors.lastname: Swanson
authors.affiliation: Department of Ecology Evolution and Behavior University of Minnesota Saint Paul MN USA.
authors.forename: Todd
authors.initials: T
authors.lastname: Arnold
authors.affiliation: Department of Ecology Evolution and Behavior University of Minnesota Saint Paul MN USA; Department of Organismic and Evolutionary Biology Harvard University Cambridge MA USA.
authors.forename: Margaret
authors.identifier: 0000-0002-1446-4856
authors.initials: M
authors.lastname: Kosmala
authors.affiliation: Department of Ecology Evolution and Behavior University of Minnesota Saint Paul MN USA.
authors.forename: James
authors.identifier: 0000-0002-5392-9556
authors.initials: J
authors.lastname: Forester
authors.affiliation: Department of Ecology Evolution and Behavior University of Minnesota Saint Paul MN USA.
authors.forename: Craig
authors.initials: C
authors.lastname: Packer
keywords: Acinonyx jubatus; Crocuta crocuta; Panthera leo; carnivore coexistence; interference competition