Spatial and temporal changes in group dynamics and range use enable anti‐predator responses in African buffalo

The views of visitors to national parks provide an important source of information to guide park planners and managers. A visitor questionnaire study was conducted in 2004 and 2005 in the Addo Elephant National Park, South Africa. A profile of the respondents, as well as records of their views on a number of park-related conservation and management issues, and of their observations of large and charismatic species (the ‘Big Five’) was compiled. The study clearly showed that national parks are important destinations for an African wildlife experience, along with other natural attractions that they have to offer. It also highlighted the scope for further visitor education, specifically aimed at improving the quality of their visits to the park. The relationship between elephant (Loxodonta africana) density and level of viewing success by visitors requires further investigation. Since the reintroduction of large predators, especially lion (Panthera leo), in the early 2000s, the daytime sighting rate of buffalo (Syncerus caffer) by visitors has increased markedly. Ongoing surveys, to monitor and expand on some of the aspects addressed in this study, are considered necessary in order to assist park staff to achieve conservation and management goals.

The mesopredator release hypothesis states that when middle-ranked predators are freed from the top-down control of apex predators, their populations will increase and predation pressure on certain prey species intensifies (Prugh et al., 2009). Examples include the release of coyotes (Canis latrans, Say 1823) following the extirpation of wolves (Canis lupus, L. 1758) in Wyoming, USA (Berger & Conner, 2008) and the suppressing effect of dingoes (Canis lupus dingo, Meyer 1793) on red fox (Vulpes vulpes, L. 1758) populations in eastern Australia (Johnson & Vanderwal, 2009). The hypothesis may provide a theoretical framework to explain the overall success of black-backed jackals (Canis mesomelas, Schreber 1775) in South Africa (Loveridge & Nel, 2004), following the extirpation of large (apex) predators, such as lions (Panthera leo, L. 1758), from many parts of their range, in the early twentieth century (Crooks & Soule, 1999). However, the increased potential for earning foreign income from trophy hunting and ⁄ or ecotourism has seen a significant change in the land use of the Eastern Cape of South Africa (Smith & Wilson, 2002). Many agrarian properties have been converted into game farms and wildlife reserves (Smith & Wilson, 2002), and large predators have been reintroduced to attract tourists (Hayward et al., 2007). The reintroduction of superior predators to game reserves may therefore reinstate top-down forcing in these systems (Estes et al., 2011), resulting in mesopredator suppression effects, such as spatial avoidance and behavioural suppression via altered activity patterns and dietary avoidance (Hayward & Kerley, 2008; Hayward & Slotow, 2009; Van Der Merwe et al., 2009). The objective of this study was to compare the dietary composition of two black-backed jackal populations; one with large predators present and one without.

We tested the widely accepted hypothesis that spotted hyaenas (Crocuta crocuta) are non-selective in their diet. The prey preference of spotted hyaena was studied in the Addo Elephant National Park (AENP), South Africa. Diet (frequency of occurrence of prey items in the diet) was quantified through the analysis of 55 scats, and compared with available prey. A combination of large- and medium-sized mammals (buffalo (Syncerus caffer), red hartebeest (Alcelaphus buselaphus) and common duiker (Sylvicapra grimmia) were the most preferred prey items. The most abundant species, warthog (Phacochoerus africanus) and kudu (Tragelaphus strepsiceros), were ignored and avoided, respectively. These results show that the assumption that hyaena prey on the most abundant available prey species may be overly simplistic. Predation patterns, such as the ones observed in AENP, may have important ramifications for less common species that are selected by hyaena in small enclosed reserves.

The reduced cost of next-generation sequencing (NGS) has allowed researchers to generate nuclear and mitochondrial genome data to gain deeper insights into the phylogeography, evolutionary history and biology of non-model species. While the Cape buffalo (Syncerus caffer caffer) has been well-studied across its range with traditional genetic markers over the last 25 years, researchers are building on this knowledge by generating whole genome, population-level data sets to improve understanding of the genetic composition and evolutionary history of the species. Using publicly available NGS data, we assembled 40 Cape buffalo mitochondrial genomes (mitogenomes) from four protected areas in South Africa, expanding the geographical range and almost doubling the number of mitogenomes available for this species. Coverage of the mitogenomes ranged from 154 to 1036X. Haplotype and nucleotide diversity for Kruger National Park (n = 15) and Mokala National Park (n = 5) were similar to diversity levels in southern and eastern Africa. Hluhluwe-iMfolozi Park (n = 15) had low levels of genetic diversity, with only four haplotypes detected, reflecting its past bottleneck. Addo Elephant National Park (n = 5) had the highest nucleotide diversity of all populations across Africa, which was unexpected, as it is known to have low nuclear diversity. This diversity was driven by a highly divergent mitogenome from one sample, which was subsequently identified in another sample via Sanger sequencing of the cytochrome b gene. Using a fossil-calibrated phylogenetic analysis, we estimated that this lineage diverged from all other Cape buffalo lineages approximately 2.51 million years ago. We discuss several potential sources of this mitogenome but propose that it most likely originated through introgressive hybridisation with an extinct buffalo species, either S. acoelotus or S. antiquus. We conclude by discussing the conservation consequences of this finding for the Addo Elephant National Park population, proposing careful genetic management to prevent inbreeding depression while maintaining this highly unique diversity.

Africa's large predator guild competes for a limited food resource base. To minimize the degree of competition, we hypothesized that the two largest members of this guild and its fiercest competitors, the lion and the spotted hyaena, would partition their activity patterns to avoid interacting. We used 96‐h continuous follows of focal animal(s) to determine when the six radio‐collared lions and eight radio‐collared spotted hyaenas, reintroduced into Addo Elephant National Park in 2003/2004, were active using a binomial measure of activity which was defined as movements >100 m during each hourly period. Contrary to our predictions, lions and hyaenas did not partition their activity times, probably because of their current low population densities. Both species exhibited a crepuscular activity pattern although hyaenas were far less active during daylight. A sub‐adult lioness minimized competitive interactions by becoming diurnal. This is likely to be a common strategy for lions that have been expelled from their natal pride to become nomadic, as it allows them to minimize kleptoparasitic and agonistic interactions from competitively dominant conspecifics and competitors. The increase in testosterone that occurs in males upon reaching sexual maturity, darkens their pelage and causes them to be more directly impacted by the heat, and thereby affords females an opportunity to escape from males during hot temperatures. Similarly, the longer pelage of young hyaenas restricts their activity to the cooler night‐time.

Simple SummaryLivestock herding is an essential and time-honored practice in Balochistan, playing a pivotal role in the region’s economy, culture, and way of life. The livestock sector is of utmost important in the province, catering to nearly 20% of the national stock. Over centuries, large predators and their prey species, including livestock, have coexisted in these mountainous landscapes. Consequently, large carnivores are more likely to interact with humans due to their extensive home ranges. This research aims to explore the impact of livestock depredation by large predators on livelihoods and conservation efforts in two districts of Balochistan, Pakistan. A human–carnivore conflict survey was conducted from July to September 2019, gathering data from 311 residents in the selected study area. Large predators in the region preyed on a total of 876 livestock over a year, comprising 560 goats, 292 sheep, 19 cows, and 5 donkeys. The gray wolf emerged as the primary predator, accounting for 66.3% of livestock depredation, which was followed by the caracal (24.3%), Asiatic jackal (8.9%), and striped hyena (0.6%). The economic loss totaled USD 78,694. Notably, 80% of respondents held negative perceptions toward wolves compared to 24.4% for caracals. Only 20.6% of respondents were aware of the importance of conserving carnivores. Livestock depredation by carnivores has fostered negative perceptions among locals toward these animals. There exists a lack of awareness regarding the significance of conserving carnivore species and their ecological roles. It is crucial to raise awareness among communities about the ecological importance of predators like the gray wolf, caracal, Asiatic jackal, striped hyena, and Balochistan black bear through community meetings and educational seminars. Furthermore, providing basic education to herders on effective livestock guarding practices is recommended to mitigate human–carnivore conflicts and promote coexistence between wildlife and local communities in Balochistan.Livestock herding is a vital practice in Balochistan, contributing to the economy and culture. The livestock sector is significant in Balochistan, providing 20% of the national stock. Large predators and their prey species, including livestock, have coexisted in these mountainous landscapes for centuries. The aim of the present research is to investigate the impacts of livestock depredation by large predators on livelihoods and predator conservation in two districts of Balochistan, Pakistan. A human–carnivore conflict survey was conducted from July to September 2019, collecting data from 311 residents in a selected study area. Large predators in the study area preyed on a total of 876 livestock during a one-year period, including 560 goats, 292 sheep, 19 cows, and 5 donkeys. The gray wolf is the leading predator, responsible for 66.3% of livestock depredation, followed by the caracal (24.3%), Asiatic jackal (8.9%), and striped hyena (0.6%). The total economic loss was USD 78,694. Overall, 80% of respondents had a negative perception of wolves compared to 24.4% for caracals. Only 20.6% of respondents knew about the importance of conserving carnivores. Livestock depredation by carnivores in the study area created a negative perception of these animals among people. There is a lack of awareness about the importance of conserving carnivore species and their role in the ecosystem. This lack of understanding has ultimately led to detrimental effects on predator populations. It is imperative to raise awareness among people about the ecological significance of carnivores through community meetings, seminars in educational institutions, and providing basic education to herders about effective livestock guarding practices.

Conservation conundrum: Endangered predators eating endangered prey

Movement patterns of lions (Panthera leo) reveal how they hunt large herbivores in heterogeneous landscapes such as the Kruger National Park in South Africa. Large herbivores are distributed differently on the landscape and therefore have different vulnerabilities as prey for lions. For instance, blue wildebeest (Connochaetes taurinus) occupy small grazing lawns at night but are difficult for lions to capture because open areas lack cover for stalking. African buffalo (Syncerus caffer) aggregate in large herds but are less available because these herds only intermittently enter the home ranges of individual lion prides. Unlike large herds of wildebeest and buffalo, plains zebra (Equus quagga) move widely in small herds while browsing greater kudus (Tragelaphus strepsiceros) and giraffes (Giraffa camelopardalis giraffa) generally occur in lower densities. We used spatial data derived from GPS collars to investigate several hypotheses regarding the movements of three lion prides in response to their prey. We found that lions were most active and moved longer distances during nighttime than during daytime. Lions remained within their core home ranges on 87% of nights and wandered to the outlying areas of the home ranges every second night. Lions visited grazing lawns, that is, area of short grass, where wildebeest herds resided every second night, and moved toward the direction of buffalo herds within 2 km of vicinity. Lions spent more time near riverbanks that provided dense woody cover at night than expected but concentrated only weakly near sites with surface water where herbivores drank in the dry season. Our study contributes to understanding how lions vary their movements in response to the spatial and temporal heterogeneity in the relative availability and vulnerability of multiple prey species.

Predation risk from lions (Panthera leo) has been linked to habitat characteristics and availability and traits of prey. We separated the effects of vegetation density and the presence of drinking water by analyzing locations of lion kills in relation to rivers with dense vegetation, which offer good lion stalking opportunities, and artificial water points with low vegetation density. The spatial distribution of lion kills was studied at the Klaserie Private Nature Reserve, South Africa. The distance between 215 lion kills and the nearest water source was analyzed using generalized linear models. Lions selected medium-sized prey species. Lion kills were closer to rivers and to artificial water points than expected by random distribution of the kills. Water that attracted prey, and not the vegetation density in riverine areas, increased predation risk, with kills of buffalo (Syncerus caffer), kudu (Tragelaphus strepsiceros), and wildebeest (Connochaetes taurinus) as water-dependent prey species. Traits of prey species, including feeding type (food habits), digestion type (ruminant or nonruminant); or body size, did not explain locations of lion kills, and no seasonal patterns in lion kills were apparent. We argue that the cascading impact of lions on local mammal assemblages is spatially heterogeneous.

Wildlife ranching, although not considered a conventional conservation system, provides a sustainable model for wildlife utilization and could be a source of valuable genetic material. However, increased fragmentation and intensive management may threaten the evolutionary potential and conservation value of species. Disease-free Cape buffalo (Syncerus caffer caffer) in southern Africa exist in populations with a variety of histories and management practices. We compared the genetic diversity of buffalo in national parks to private ranches and found that, except for Addo Elephant National Park, genetic diversity was high and statistically equivalent. We found that relatedness and inbreeding levels were not substantially different between ranched populations and those in national parks, indicating that breeding practices likely did not yet influence genetic diversity of buffalo on private ranches in this study. High genetic differentiation between South African protected areas highlighted their fragmented nature. Structure analysis revealed private ranches comprised three gene pools, with origins from Addo Elephant National Park, Kruger National Park and a third, unsampled gene pool. Based on these results, we recommend the Addo population be supplemented with disease-free Graspan and Mokala buffalo (of Kruger origin). We highlight the need for more research to characterize the genetic diversity and composition of ranched wildlife species, in conjunction with wildlife ranchers and conservation authorities, in order to evaluate the implications for management and conservation of these species across different systems.

Wildlife ranching, although not considered a conventional conservation system, provides a sustainable model for wildlife utilization and could be a source of valuable genetic material. However, increased fragmentation and intensive management may threaten the evolutionary potential and conservation value of species. Disease-free Cape buffalo (Syncerus caffer caffer) in southern Africa exist in populations with a variety of histories and management practices. We compared the genetic diversity of buffalo in national parks to private ranches and found that, except for Addo Elephant National Park, genetic diversity was high and statistically equivalent. We found that relatedness and inbreeding levels were not substantially different between ranched populations and those in national parks, indicating that breeding practices likely did not yet influence genetic diversity of buffalo on private ranches in this study. High genetic differentiation between South African protected areas highlighted their fragmented nature. Structure analysis revealed private ranches comprised three gene pools, with origins from Addo Elephant National Park, Kruger National Park and a third, unsampled gene pool. Based on these results, we recommend the Addo population be supplemented with disease-free Graspan and Mokala buffalo (of Kruger origin). We highlight the need for more research to characterize the genetic diversity and composition of ranched wildlife species, in conjunction with wildlife ranchers and conservation authorities, in order to evaluate the implications for management and conservation of these species across different systems.

Despite extensive evidence that African buffalo Syncerus caffer are grazers, De Graaff et al. using rumen content analysis of animals that had starved to death proposed that buffalo in grass-limited Eastern Cape thicket should be considered browsers. Although these anomalous findings were initially accepted, but later challenged, the browse-dominated diet continues to be used as a foundation for hypotheses on the diet of healthy animals. Consequently, the debate around buffalo as browsers or grazers in thicket has not yet been settled. We describe the diet of buffalo in the Addo Elephant National Park and include data from other published work from the region to test the importance of grass in buffalo diet. We show that the diet is dominated by grasses, even in grass-limited thicket, and that browse species are seldom dominant foods. Thus, there is no empirical evidence to corroborate the notion that buffalo switch their diet to browse when grass availability is low. In an attempt to advance our understanding of buffalo foraging in thicket, we reiterate that De Graaff’s work is not a valid measure of buffalo diet in succulent thicket and that additional testing of the browser–grazer hypothesis is not needed.Conservation implications: Our results confirm that buffalo are grazers, rather than browsers, in grass-limited Eastern Cape thicket. Thus, additional testing of the browser–grazer hypothesis for buffalo in the region is not needed.

Elephants facilitate impact of large predators on small ungulate prey species

Management interventions in small (<30 000 ha) to medium-sized (<100 000 ha) fenced reserves require careful planning to minimize alterations to predator-prey dynamics. However, when interventions result in severe changes to predator prey dynamics, learning from these outcomes can assist in informing future interventions. In small to medium-sized ventions. We used lion (Panthera leo) and buffalo (Syncerus caffer) interventions (lion introduction and buffalo translocation) in the Addo Elephant National Park to investigate the response of the buffalo—lion dynamic in the park. Responses did not conform to our expectations for either lions or buffalo. Rather than dominate the remaining solitary male lion, the introduced coalition split, with one of the introduced male lions (known to hunt buffalo successfully where it originated) joining with the remaining solitary male. Following fragmentation as a result of the buffalo translocation, previously large buffalo herds failed to re-form. These two outcomes resulted in a significant increase in buffalo predation primarily by the newly formed coalition of males, with the majority of buffalo killed originating from splintered herds rather than male-dominated groups as is conventionally found in large ecosystems. Ultimately, management actions can enable us to learn by monitoring the consequences of interventions, allowing us to implement what we have learned in an adaptive management framework.

1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.