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 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.
Highlights
| METHODSOur 1,125-km study area (bounded in the northwest at lat/long of −2.363589, 34.72594; in the southeast at −2.660651, 35.18051) was located at the center of Serengeti National Park, Tanzania, at the intersection of open plains and savanna woodlands (Figure 1a)
We evaluated the relative importance of intraguild predators using analysis of deviance (ANODEV; Harris et al, 2005) and comparing models with and without covariates indicating the presence and (a)
Subordinate competitors are expected to seek out “competition refuges” by selecting marginal habitats (Durant, 1998; Linnell & Strand, 2000), we found that interference competition among lions, hyenas, and cheetahs did not translate into long-term displacement by subordinate species, even at fine spatial scales
Summary
Our 1,125-km study area (bounded in the northwest at lat/long of −2.363589, 34.72594; in the southeast at −2.660651, 35.18051) was located at the center of Serengeti National Park, Tanzania, at the intersection of open plains and savanna woodlands (Figure 1a). We evaluated patterns of fine-scale spatial avoidance among lions, hyenas, and cheetahs by comparing species-specific capture rates at each site. We calculated expected values for number of observations during each 12-hr bin using a discrete time-to-event model that assumed exponential decay in the probability that the species of interest had not been seen yet, with expected encounter probabilities calculated using pooled information on detection probability throughout the first 6 days, with an adjustment based on probability of activity during each 12-hr bin based on diel activity patterns for the second observed species This assured that observed patterns were not due to similar or contrasting diel patterns (e.g., a strictly nocturnal species not observed in the 12 hr following an early morning observation of a mostly diurnal species might reflect temporal niche partitioning rather than active avoidance; see Swanson, 2014). Whereas lion sightings remained significantly higher for 24 hr following hyena sightings, hyena sightings declined sharply following 12 hr after a lion sighting (Table 3; Figure 6)
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