Spatial heterogeneity facilitates carnivore coexistence.

Competitively dominant carnivore species can limit the population sizes and alter the behavior of inferior competitors. Established mechanisms that enable carnivore coexistence include spatial and temporal avoidance of dominant predator species by subordinates, and dietary niche separation. However, spatial heterogeneity across landscapes could provide inferior competitors with refuges in the form of areas with lower competitor density and/or locations that provide concealment from competitors. Here, we combine temporally overlapping telemetry data from dominant lions (Panthera leo) and subordinate African wild dogs (Lycaon pictus) with high-resolution remote sensing in an integrated step selection analysis to investigate how fine-scaled landscape heterogeneity might facilitate carnivore coexistence in South Africa's Hluhluwe-iMfolozi Park, where both predators occur at exceptionally high densities. We ask whether the primary lion-avoidance strategy of wild dogs is spatial avoidance of lions or areas frequented by lions, or if wild dogs selectively use landscape features to avoid detection by lions. Within this framework, we also test whether wild dogs rely on proactive or reactive responses to lion risk. In contrast to previous studies finding strong spatial avoidance of lions by wild dogs, we found that the primary wild dog lion-avoidance strategy was to select landscape features that aid in avoidance of lion detection. This habitat selection was routinely used by wild dogs, and especially when in areas and during times of high lion-encounter risk, suggesting a proactive response to lion risk. Our findings suggest that spatial landscape heterogeneity could represent an alternative mechanism for carnivore coexistence, especially as ever-shrinking carnivore ranges force inferior competitors into increased contact with dominant species.