Most ecosystems have multiple predator species that not only compete for shared prey, but also pose direct threats to each other. These intraguild interactions are key drivers of carnivore community structure, with ecosystem-wide cascading effects. Yet, behavioral mechanisms for coexistence of multiple carnivore species remain poorly understood. The challenges of studying large, free-ranging carnivores have resulted in mainly coarse-scale examination of behavioral strategies without information about all interacting competitors. We overcame some of these challenges by examining the concurrent fine-scale movement decisions of almost all individuals of four large mammalian carnivore species in a closed terrestrial system. We found that the intensity ofintraguild interactions did not follow a simple hierarchical allometric pattern, because spatial and behavioral tactics of subordinate species changed with threat and resource levels across seasons. Lions (Panthera leo) were generally unrestricted and anchored themselves in areas rich in not only their principal prey, but also, during periods of resource limitation (dry season), rich in the main prey for other carnivores. Because of this, the greatest cost (potential intraguild predation) for subordinate carnivores was spatially coupled with the highest potential benefit of resource acquisition (prey-rich areas), especially in the dry season. Leopard (P. pardus) and cheetah (Acinonyx jubatus) overlapped with the home range of lions but minimized their risk using fine-scaled avoidance behaviors and restricted resource acquisition tactics. The cost of intraguild competition was most apparent for cheetahs, especially during the wet season, as areas with energetically rewarding large prey (wildebeest) were avoided when they overlapped highly with the activity areas of lions. Contrary to expectation, the smallest species (African wild dog, Lycaon pictus) did not avoid only lions, but also used multiple tactics to minimize encountering all other competitors. Intraguild competition thus forced wild dogs into areas with the lowest resource availability year round. Coexistence of multiple carnivore species has typically been explained by dietary niche separation, but our multi-scaled movement results suggest that differences in resource acquisition may instead be a consequence of avoiding intraguild competition. We generate a more realistic representation of hierarchical behavioral interactions that may ultimately drive spatially explicit trophic structures of multi-predator communities.

Studies that focus on single predator-prey interactions can be inadequate for understanding antipredator responses in multi-predator systems. Yet there is still a general lack of information about the strategies of prey to minimize predation risk from multiple predators at the landscape level. Here we examined the distribution of seven African ungulate species in the fenced Karongwe Game Reserve (KGR), South Africa, as a function of predation risk from all large carnivore species (lion, leopard, cheetah, African wild dog, and spotted hyena). Using observed kill data, we generated ungulate-specific predictions of relative predation risk and of riskiness of habitats. To determine how ungulates minimize predation risk at the landscape level, we explicitly tested five hypotheses consisting of strategies that reduce the probability of encountering predators, and the probability of being killed. All ungulate species avoided risky habitats, and most selected safer habitats, thus reducing their probability of being killed. To reduce the probability of encountering predators, most of the smaller prey species (impala, warthog, waterbuck, kudu) avoided the space use of all predators, while the larger species (wildebeest, zebra, giraffe) only avoided areas where lion and leopard space use were high. The strength of avoidance for the space use of predators generally did not correspond to the relative predation threat from those predators. Instead, ungulates used a simpler behavioral rule of avoiding the activity areas of sit-and-pursue predators (lion and leopard), but not those of cursorial predators (cheetah and African wild dog). In general, selection and avoidance of habitats was stronger than avoidance of the predator activity areas. We expect similar decision rules to drive the distribution pattern of ungulates in other African savannas and in other multi-predator systems, especially where predators differ in their hunting modes.

Summary 1.We evaluated one of the most extensive efforts to date to re-introduce an endangered species: attempts to establish an actively managed meta-population of African wild dogs Lycaon pictus in South Africa. 2. Using an information-theoretic approach, known-fate modelling in program mark was employed to estimate the survival of re-introduced wild dogs and their offspring, and to model covariate effects relative to survival. Multiple a priori hypotheses on correlates of re-introduction success were tested (collated from extensive individual experiences) using different re-introduction attempts as natural quasi experiments. 3. Survival analyses revealed that the determinants of re-introduction success can be reduced to two factors relevant for management, suggesting that wild dog re-introductions should be attempted with socially integrated animals that are released into securely fenced areas, unless measures are implemented to mitigate human-related mortalities outside protected areas. 4. Synthesis and application. This study illustrates that monitoring and evaluation of conservation efforts, complimented with expert knowledge, forms the foundation of informed decision-making to underpin management recommendations with scientific evidence, particularly if the proposed actions are controversial.

Estimation of survival rates is important for developing and evaluating conservation options for large carnivores. However, telemetry studies for large carnivores are often characterized by small sample sizes that limit meaningful conclusions. We used data from 10 published and 8 unpublished studies of leopards Panthera pardus in southern Africa to estimate survival rates and investigate causes of leopard mortality. Mean survival rates were significantly lower in non-protected (0.55 ± SE 0.08) compared to protected areas (0.88 ± 0.03). Inside protected areas juveniles had significantly lower survival (0.39 ± 0.10) compared to subadults (0.86 ± 0.07) and adults (0.88 ± 0.04).There was a greater difference in cause of death between protected and non-protected areas for females compared to males, with people being the dominant cause of mortality outside protected areas for both females and males. We suggest there is cause for concern regarding the sustainability of leopard populations in South Africa, as high female mortality may have severe demographic effects and a large proportion of suitable leopard habitat lies in non-protected areas. However, because a large proportion of deaths outside protected areas were attributed to deliberate killing by people, we suggest that management interventions may have the potential to increase leopard survival dramatically. We therefore stress the urgency to initiate actions, such as conflict mitigation programmes, to increase leopard survival in non-protected areas.