In the absence of a “landscape of fear”: How lions, hyenas, and cheetahs coexist

Alexandra Swanson, Todd Arnold, Margaret Kosmala, James Forester, Craig Packer

Research output: Contribution to journalArticlepeer-review

40 Scopus citations


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 km2 in 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.

Original languageEnglish (US)
Pages (from-to)8534-8545
Number of pages12
JournalEcology and Evolution
Issue number23
StatePublished - Dec 1 2016

Bibliographical note

Funding Information:
Research clearance was provided by the Tanzania Wildlife Research Institute and Tanzania National Parks. We thank members of the Serengeti Lion Project and the 28,040 volunteers who contributed to Snapshot Serengeti classifications (complete list at This work was supported by National Science Foundation grant DEB-1020479 to CP for maintenance of the long-term Lion Project, the University of Minnesota Supercomputing Institute, private donations raised during the Serengeti Live and Save Snapshot Serengeti crowd-funding campaigns, and by grants to AS from Explorer's Club, UMN Thesis Research Grants, UMN Office of International Programs, American Society of Mammalogists, and Minnesota Zoo Ulysses S. Seal Conservation fund. Snapshot Serengeti website development was funded by awards to the Zooniverse from the Alfred P. Sloan Foundation.

Publisher Copyright:
© 2016 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.


  • Acinonyx jubatus
  • Crocuta crocuta
  • Panthera leo
  • carnivore coexistence
  • interference competition


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