The size of the home range of a mammal is affected by numerous factors. However, in the normally solitary, but polygynous, Leopard (Panthera pardus), home range size and maintenance is complicated by their transitory social grouping behavior, which is dependent on life history stage and/or reproductive status. In addition, the necessity to avoid competition with conspecifics and other large predators (including humans) also impacts upon home range size. We used movement data from 31 sites across Africa, comprising 147 individuals (67 males and 80 females) to estimate the home range sizes of leopards. We found that leopards with larger home ranges, and in areas with more vegetation, spent longer being active and generally traveled faster, and in straighter lines, than leopards with smaller home ranges. We suggest that a combination of bottom-up (i.e., preferred prey availability), top-down (i.e., competition with conspecifics), and reproductive (i.e., access to mates) factors likely drive the variability in Leopard home range sizes across Africa. However, the maintenance of a large home range is energetically expensive for leopards, likely resulting in a complex evolutionary trade-off between the satisfaction of basic requirements and preventing potentially dangerous encounters with conspecifics, other predators, and people. Bringing all the boys to the yard. Male leopards travel further and faster to find females and to avoid other males. Whereas females are more concerned with food than other leopards.

Spatial patterns of and competition for resources by territorial carnivores are typically explained by two hypotheses: 1) the territorial defence hypothesis and 2) the searching efficiency hypothesis. According to the territorial defence hypothesis, when food resources are abundant, carnivore densities will be high and home ranges small. In addition, carnivores can maximise their necessary energy intake with minimal territorial defence. At medium resource levels, larger ranges will be needed, and it will become more economically beneficial to defend resources against a lower density of competitors. At low resource levels, carnivore densities will be low and home ranges large, but resources will be too scarce to make it beneficial to defend such large territories. Thus, home range overlap will be minimal at intermediate carnivore densities. According to the searching efficiency hypothesis, there is a cost to knowing a home range. Larger areas are harder to learn and easier to forget, so carnivores constantly need to keep their cognitive map updated by regularly revisiting parts of their home ranges. Consequently, when resources are scarce, carnivores require larger home ranges to acquire sufficient food. These larger home ranges lead to more overlap among individuals' ranges, so that overlap in home ranges is largest when food availability is the lowest. Since conspecific density is low when food availability is low, this hypothesis predicts that overlap is largest when densities are the lowest. We measured home range overlap and used a novel method to compare intraspecific home range overlaps for lions Panthera leo (n = 149) and leopards Panthera pardus (n = 111) in Africa. We estimated home range sizes from telemetry location data and gathered carnivore density data from the literature. Our results did not support the territorial defence hypothesis for either species. Lion prides increased their home range overlap at conspecific lower densities whereas leopards did not. Lion pride changes in overlap were primarily due to increases in group size at lower densities. By contrast, the unique dispersal strategies of leopards led to reduced overlap at lower densities. However, when human-caused mortality was higher, leopards increased their home range overlap. Although lions and leopards are territorial, their territorial behaviour was less important than the acquisition of food in determining their space use. Such information is crucial for the future conservation of these two iconic African carnivores.

Home range size is a fundamental measure of animal space use, providing insight into habitat quality, animal density, and social organization. Human impacts increasingly are affecting wildlife, especially among wide-ranging species that encounter anthropogenic disturbance. Leopards (Panthera pardus) provide a useful model for studying this relationship because leopards coexist with people at high and low human densities and are sensitive to human disturbance. To compare leopard home range size across a range of human densities and other environmental conditions, we combined animal tracking data from 74 leopards in multiple studies with new analytical techniques that accommodate different sampling regimes. We predicted that home ranges would be smaller in more productive habitats and areas of higher human population density due to possible linkage with leopard prey subsidies from domestic species. We also predicted that male leopards would have larger home ranges than those of females. Home ranges varied in size from 14.5 km2 in India to 885.6 km2 in Namibia, representing a 60-fold magnitude of variation. Home range stability was evident for 95.2% of nontranslocated individuals and 38.5% of translocated individuals. Leopard home range sizes were negatively correlated with landscape productivity, and males used larger areas than females. Leopards in open habitats had a predicted negative correlation in home range size with human population density, but leopards in closed habitats used larger home ranges in areas with more people.

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 sig- nificantly 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) com- pared 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 sustain- ability 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 pro- portion of deaths outside protected areas were attributed to deliberate killing by people, we suggest that manage- ment 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.

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.

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.

Background: Group dynamics of gregarious ungulates in the grasslands of the African savanna have been well studied, but the trade-offs that affect grouping of these ungulates in woodland habitats or dense vegetation are less well understood. We examined the landscape-level distribution of groups of blue wildebeest, Connochaetes taurinus, and Burchell's zebra, Equus burchelli, in a predominantly woodland area (Karongwe Game Reserve, South Africa; KGR) to test the hypothesis that group dynamics are a function of minimizing predation risk from their primary predator, lion, Panthera leo.Methodology/Principal Findings: Using generalized linear models, we examined the relative importance of habitat type (differing in vegetation density), probability of encountering lion (based on utilization distribution of all individual lions in the reserve), and season in predicting group size and composition. We found that only in open scrub habitat, group size for both ungulate species increased with the probability of encountering lion. Group composition differed between the two species and was driven by habitat selection as well as predation risk. For both species, composition of groups was, however, dominated by males in open scrub habitats, irrespective of the probability of encountering lion.Conclusions/Significance: Distribution patterns of wildebeest and zebra groups at the landscape level directly support the theoretical and empirical evidence from a range of taxa predicting that grouping is favored in open habitats and when predation risk is high. Group composition reflected species-specific social, physiological and foraging constraints, as well as the importance of predation risk. Avoidance of high resource open scrub habitat by females can lead to loss of foraging opportunities, which can be particularly costly in areas such as KGR, where this resource is limited. Thus, landscape-level grouping dynamics are species specific and particular to the composition of the group, arising from a tradeoff between maximizing resource selection and minimizing predation risk.

Leopards (Panthera pardus) are a poorly understood, solitary felid, and additional study could provide novel insights into both evolution and conservation management. We studied the reproductive biology of 7 adult female and 2 adult male leopards on Karongwe, South Africa, from 1999 to 2005. We intensively researched copulatory biology from 2001 to 2003, during which we observed 19 consortships and 4,855 copulations (1,809 copulations visually) during 2,449 h of following consort pairs. Leopards copulated on average 4 times per hour, with an average of 256 copulations per consortship. Conception rate was low, resulting, on average, after 2.3 consortships (SE = 0.4 consortships). All reproductive parameters except gestation length were shorter than those in published literature, which we suspect reflected a facultative response to superabundant resources. Although females produced the expected 1.6 cubs per year, lifetime reproductive rates on Karongwe Reserve were approximately half that reported elsewhere due to lower female survival resulting from intraspecific factors. These results offer encouragement for founding new populations of endangered felids in areas of high prey availability and resource abundance, provided factors affecting population regulation can be managed. DOI: 10.1644/09-MAMM-A-256.1.

The prey selection and predatory behaviour of a single pride of lions (Panthera leo) was studied in the 8500 ha Karongwe Game Reserve from 1999–2005. The study focused on the difference between prey selection in the first three years when a two-male coalition was present with a similar period thereafter when one of the males was removed and subadult males dispersed from the pride. A total of 662 kills were recorded, with blue wildebeest (Connochaetes taurinus), warthog (Phacochoerus africanus), waterbuck (Kobus ellipsiprymnus), zebra (Equus burchelli) and impala (Aepyceros melampus) being the most preferred species. Although there was preference for these species, and adult prey were favoured, there was no significant selection for prey size, age or gender. There was a significant difference in the number of kills made when the two-male coalition was present as opposed to the single pride male. The solitary pride male spent significantly more time with the females and shared most of their kills, and during this period dispersing subadult males killed a substantial amount of prey when separated from the pride. Knowledge on predator feeding behaviour is important for managers to assess impact on prey populations in small reserves in order to manage them correctly, thereby preventing depletion of such populations.

The reproductive biology of a pride of lions (Panthera leo) was studied on the 8500 ha Karongwe Game Reserve from 1999 to 2005. Over this period, the pride consisted of between four and 11 lions with a paired coalition of adult males during the first three years and a single adult male for the next three years. We recorded shorter than normal interbirth intervals, high birth rates of 1.3 cubs/lioness/year, very high cub survival rates, and subadults leaving the pride at young ages. This translated into substantially faster growth rates than are typical in large lion populations in ecologically similar circumstances such as Kruger National Park, but are similar to those of lions in Serengeti National Park. These demographic characteristics were probably induced initially by a lack of intense intraspecific competition and high prey availability, but population stability was maintained through the removal of young subadults by management. Interestingly, variability in conception rates between lionesses resulted in lower growth rates than have been found in other similar reintroduced lion populations. These demographic traits characterize many of the small reintroduced lion populations, and call for appropriate management to avoid the consequences of high predator densities.

Summary 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. 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. 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. 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.

Interventionist conservation management of territorial large carnivores has increased in recent years, especially in South Africa. Understanding of spatial ecology is an important component of predator conservation and management. Spatial patterns are influenced by many, often interacting, factors making elucidation of key drivers difficult. We had the opportunity to study a simplified system, a single pride of lions (Panthera leo) after reintroduction onto the 85 km(2) Karongwe Game Reserve, from 1999-2005, using radio-telemetry. In 2002 one male was removed from the paired coalition which had been present for the first three years. A second pride and male were in a fenced reserve adjacent of them to the east. This made it possible to separate social and resource factors in both a coalition and single male scenario, and the driving factors these seem to have on spatial ecology. Male ranging behaviour was not affected by coalition size, being driven more by resource rather than social factors. The females responded to the lions on the adjacent reserve by avoiding the area closest to them, therefore females may be more driven by social factors. Home range size and the resource response to water are important factors to consider when reintroducing lions to a small reserve, and it is hoped that these findings lead to other similar studies which will contribute to sound decisions regarding the management of lions on small reserves.