Large carnivore populations are declining worldwide due to anthropogenic causes such as habitat loss and human expansion into wild areas. Competition between large carnivores can exacerbate this decline. While brown hyena Parahyaena brunnea and spotted hyena Crocuta crocuta belong to the same family, they are rarely found in the same area or co-occur at low densities as spotted hyena are known to exclude brown hyena. In Central Tuli, Botswana, however, brown hyena and spotted hyena are both found at high densities. We undertook a camera trap survey in this area to estimate the densities of both species, and to examine temporal overlap and co-detection patterns of brown and spotted hyena. Estimated population densities based on spatial capture-recapture models were 10.5 AE 1.9/100 km 2 for brown hyena and 14.9 AE 2.2/100 km 2 for spotted hyena. These population densities are among the highest reported estimates in southern Africa. Strong temporal overlap was found between brown and spotted hyena, while there was no decrease in detection rate of brown hyena at camera sites where spotted hyena were also detected, which indicates that both hyena species did not tend to avoid encounters. Although both species compete for the same prey, we suggest as possible explanations that prey densities are high and that competition does not significantly negatively impact brown hyena, because brown hyena is a scavenger whereas spotted hyena scavenge and kill prey. With the found high densities of both carnivores, this study adds to the known variation in composition of existing large carnivore communities and suggests testable explanations for these densities.

Intraguild interactions among mammalian carnivores are important in shaping carnivore guild composition. Competing species may inhabit different areas and/or being active during different times to reduce the risk of aggressive interactions, but the role of body size in intraguild interactions within carnivore guilds remains largely unknown. We determined spatial and temporal co‐occurrence of small, medium‐sized and large carnivores of the carnivore guild in central Tuli, Botswana: lion Panthera leo, leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea, black‐backed jackal Canis mesomelas, bat‐eared fox Otocyon megalotis, African wildcat Felis sylvestris lybica, African civet Civettictis civetta, honey badger Mellivora capensis and small‐spotted genet Genetta genetta. We used camera trap data over a two‐year period and quantified the degree of temporal and spatial overlap by comparing activity patterns and calculating Pianka's index respectively. Our results showed that temporal overlap in activity between all carnivore species was high, but complete overlap was possibly reduced by differences in peak activity periods. In addition, low to moderate levels of spatial overlap were found between the different carnivore species, supporting the idea that small carnivore species inhabit different areas than large species to reduce the risk of interference competition. Due to the possible strong competition amongst sympatric carnivores there is a need for more knowledge on co‐existence patterns for successful management and conservation of carnivore species, for example when carnivore species are (re)introduced in an area.

Scavenging is widespread in the carnivore guild and can greatly impact food web structures and population dynamics by either facilitation or suppression of sympatric carnivores. Due to habitat loss and fragmentation, carnivores are increasingly forced into close sympatry, possibly resulting in more interactions such as kleptoparasitism and competition. In this paper, we investigate the potential for these interactions when carnivore densities are high. A camera trap survey was conducted in central Tuli, Botswana, to examine leopard Panthera pardus densities and spatiotemporal activity patterns of leopard and its most important competitors' brown hyena Parahyaena brunnea and spotted hyena Crocuta crocuta. Spatial capture–recapture models estimated leopard population density to be 12.7 ± 3.2 leopard/100 km 2 , which is one of the highest leopard densities in Africa. Time‐to‐event analyses showed both brown hyena and spotted hyena were observed more frequently before and after a leopard observation than expected by chance. The high spatiotemporal overlap of both hyena species with leopard is possibly explained by leopard providing scavenging opportunities for brown hyena and spotted hyena. Our results suggest that central Tuli is a high‐density leopard area, despite possible intense kleptoparasitism and competition.

Large carnivore community structure is affected by direct and indirect interactions between intra‐guild members. Co‐existence between different species within a carnivore guild may occur through diet, habitat or temporal partitioning. Since carnivore species are highly dependent on availability and accessibility of prey, diet partitioning is potentially one of the most important mechanisms in allowing carnivores to co‐exist. Intra‐guild interactions may vary over time as carnivore prey preference and diet overlap can change due to seasonal changes in resource availability. We conducted scat analysis to compare the seasonal changes in prey preference, diet partitioning and niche breadth of four large carnivore species, namely leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea and wild dog Lycaon pictus in central Tuli, Botswana. Large carnivores in central Tuli display a high dietary overlap, with spotted hyena and brown hyena displaying almost complete dietary overlap and the other carnivore species displaying slightly lower but still significant dietary overlap. Dietary niche breadth for both hyena species was high possibly due to their flexible foraging strategies, including scavenging, while leopard and wild dog showed a relatively low niche breadth, suggesting a more specialised diet. High dietary overlap in central Tuli is possibly explained by the high abundance of prey species in the area thereby reducing competition pressure between carnivore species. Our research highlights the need to assess the influence of diet partitioning in structuring large carnivore communities across multiple study sites, by demonstrating that in prey rich environments, the need for diet partitioning by carnivores to avoid competition may be limited.

Table 1.1: Comparison of life-history traits and behavior between the main focal species: leopard, spotted hyena and brown hyena Leopard Panthera pardus Spotted Hyena Crocuta crocuta Brown Hyena Parahyaena brunnea Body weight -Male -Female 20 -90 kg a 17 -60 kg a 60 kg b 70 kg b No significant sexual dimorphism between males and females c Average weight is 40 kg a Litter size 1 -3 cubs a 1 -2 cubs b 1 -4 cubs a Home range size Wide variation, from 2104.4 km 2 for males and 1285.8 km 2 for females in the Kgalagadi Transfrontier Park in Southern Africa d to 32.8 km 2 for males and 14.0 km 2 for females in the Lolldaiga hills in Kenya e Wide variation, from 40 km 2 to 1000 km 2 b Wide variation, from 215km 2 to 461km 2 in the southern Kalahari f to 16 km 2 to 97 km 2 in northcentral Namibia g Population size Earlier Africa-wide assessments of population size were criticized as being too high h . The most commonly cited number is 700,000 + in Africa i although calculation was reported similarly flawed. 27000 -47000 individuals j 4365 -10111 mature individuals k Main prey species Preferentially prey upon species within a weight range of 10-40 kg, most common prey species are impala, bushbuck and common duiker l