Ecological and anthropogenic influences on space use by spotted hyaenas

Due to increasing human encroachment into the remaining habitat of many large carnivore species, there is an immediate need to understand the ecological and anthropogenic factors influencing carnivore space use decisions. In particular, knowledge of changes in space use in response to disturbance, and the costs associated with these changes, will be critical in guiding conservation efforts. To investigate the ecological factors influencing carnivore space use, we intensively radiotracked members of two large social groups (clans) of spotted hyaenas Crocuta crocuta in the Masai Mara National Reserve, Kenya. In addition, we studied the influence of livestock grazing by comparing space use between two study clans that differed dramatically in exposure to grazing. Logistic regression modeling indicated that space use in the absence of livestock was most influenced by the location of the clan's communal den. However, hyaenas were also found to select shrubland, areas of high prey density, and proximity to seasonal streams. Movements of hyaenas exposed to livestock grazing were most influenced by vegetation type, with a strong avoidance of open grass plains. Den location and prey density had less influence on space use decisions in the disturbed than the undisturbed clan. Livestock distribution did not directly influence hyaena movements either during daytime, when livestock were present, or at night. We suggest that direct livestock avoidance was unnecessary due to the observed increased use of vegetative cover by hyaenas exposed to grazing livestock. The greater distances from the den, and from areas of high prey density at which hyaenas were found in disturbed than undisturbed areas indicates potential energetic costs incurred by disturbed hyaenas. Our results therefore suggest that reduced vegetative cover, as is often found outside protected areas, may result in more dramatic modifications of hyaena movements in the presence of livestock.     

Densities of spotted hyaena (Crocuta crocuta) and African golden wolf (Canis anthus) increase with increasing anthropogenic influence

We report densities of spotted hyaena (Crocuta crocuta) and African golden wolf (Canis anthus) in Enderta district in northern Ethiopia with high human and low natural prey densities. We estimated spotted hyaena and African golden wolf abundance and characterized their spatial distribution with three methods we surveyed four road-transects for 66 nights during dry (n = 41) and wet (n = 25) seasons, we used 34 calling stations and we mapped all active spotted hyaena dens at the time of the survey. The density of spotted hyaena and African golden wolf increased with proximity to towns where human density was higher. A total of 562 spotted hyaena and 63 African golden wolf responded to calling stations, leading to estimates of 1145 spotted hyaena and 166 African golden wolf in Enderta district. This method also found a significantly higher spotted hyaena and African golden wolf abundance in high human density areas. Maximum response radius was 2.8 km for spotted hyaena and 2.5 km for African golden wolf, and response probability was 0.83 for spotted hyaena and 0.8 for African golden wolf, respectively. We found 40 active spotted hyaena dens with 1507 remnants of prey, and the majority of the dens were located close to rivers and villages. Our findings show a positive relationship between spotted hyaena, African golden wolf and human concentrations that might demonstrate a case of exceptional coexistence of humans and carnivores, both at high densities. We suggest further investigations into co-adaptations between humans and predators in the study area.     

Heterogeneity in the density of spotted hyaenas in Hluhluwe-iMfolozi Park,South Africa

Animal population sizes and trends, as well as their distributions, are essential information to the understanding and conservation of ecosystems. During this study in Hluhluwe-iMfolozi Park, South Africa, a spotted hyaena Crocuta crocuta Erxleben, 1777 (Hyaenidae) population was surveyed by attracting individuals with pre-recorded sounds. The hyaena population (excluding cubs) is substantially larger (321 individuals) than the previous estimate of 200 and this population is the second largest protected population in South Africa. Average hyaena density, at 0.357 individuals/km², was relatively high compared to other southern African conservation areas, and range from 0 to 1.25 individuals/km² across sampling stations. For short periods, spatial heterogeneity in density was marked at small and large spatial scales, but decreased when averaged over a longer period. This heterogeneity may be important in promoting the coexistence of other large and mobile carnivores in Hluhluwe-iMfolozi Park by creating potential dynamic competition refugia in space and time. Furthermore, heterogeneity of hyaena density at smaller scales should influence studies investigating the avoidance of hyaenas by competitively inferior carnivores.     

The Lion King and the Hyaena Queen: large carnivore interactions and coexistence

Interactions among species, which range from competition to facilitation, have profound effects on ecosystem functioning. Large carnivores are of particular importance in shaping community structure since they are at the top of the food chain, and many efforts are made to conserve such keystone species. Despite this, the mechanisms of carnivore interactions are far from understood, yet they are key to enabling or hindering their coexistence and hence are highly relevant for their conservation. The goal of this review is thus to provide detailed information on the extents of competition and facilitation between large carnivores and their impact in shaping their life histories. Here, we use the example of spotted hyaenas (Crocuta crocuta) and lions (Panthera leo) and provide a comprehensive knowledge of their interactions based on meta‐analyses from available literature (148 publications). Despite their strong potential for both exploitation and interference competition (range and diet overlap, intraguild predation and kleptoparasitism), we underline some mechanisms facilitating their coexistence (different prey‐age selection and scavenging opportunities). We stress the fact that prey abundance is key to their coexistence and that hyaenas forming very large groups in rich ecosystems could have a negative impact on lions. We show that the coexistence of spotted hyaenas and lions is a complex balance between competition and facilitation, and that prey availability within the ecosystem determines which predator is dominant. However, there are still many gaps in our knowledge such as the spatio‐temporal dynamics of their interactions. As both species' survival becomes increasingly dependent on protected areas, where their densities can be high, it is critical to understand their interactions to inform both reintroduction programs and protected area management.     

How much does a spotted hyaena eat? Perspective from the Namib Desert

The amount of food consumed by a clan of spotted hyaenas in the Namib desert was determined gravimetrically for 19 nights and the amount consumed by each individual was calculated according to its feeding time budget. At large, fleshy ungulate carcasses, each adult clan member ate about 8.7kg nightly, except for the bottom-ranking male, who obtained only half that amount per night. Skin and bone remains were consumed at a slower rate of 2kg/hyaena/night. Medium-sized ungulates were devoured rapidly so that each hyaena obtained more during such feeding nights than at large carcasses. Over a 29 day period, a hyaena clan consumed five cacasses at a rate of 4.0kg/hyaena/day, which compared well with previous estimates of average daily consumption in large social carnivores studied elsewhere. Annually, sixteen Namib hyaenas ate some 4.8% of the ungulate bio-mass, removing, in terms of numbers 14.3% of the gemsbok and 2.2% of the mountain zebra. This depletion did not appear to limit these populations.     

Dietary partitioning of three large carnivores in Majete Wildlife Reserve,Malawi

Between 2011 and 2012, the carnivore guild in Majete Wildlife Reserve (MWR), Malawi, was restored following the reintroduction of lion (Panthera leo) and leopard (Panthera pardus). The aim of this study was to describe and compare the diet of lion, leopard and resident spotted hyaena (Crocuta crocuta) using scat analysis. Lions and spotted hyaenas displayed the greatest dietary overlap (O_ab = 0.88) and selected mainly medium- to large-bodied prey species. Lions had a mean preferred prey weight of 120.33 ± 42.14 kg (SE), with warthog (Phacochoerus africanus) and waterbuck (Kobus ellipsiprymnus) making up 60.64% of relative biomass consumed. Spotted hyaenas had a mean preferred prey weight of 102.40 ± 41.69 kg and had a more generalised diet (B_a = 0.46) compared to lions (B_a = 0.36). In contrast, leopards occupied a dietary niche substantially lower than that of lions and spotted hyaenas, selecting relatively smaller prey with a mean preferred prey weight of 27.50 ± 6.74 kg. Our results suggest that coexistence between the resident hyaena and reintroduced lion and leopard in MWR is facilitated by dietary partitioning. We advise long-term monitoring of reintroduced carnivores in small, enclosed reserves to assess their impacts on predator and prey populations.     

Prey preferences of the spotted hyaena (Crocuta crocuta) and degree of dietary overlap with the lion (Panthera leo)

Spotted hyaenas Crocuta crocuta were once considered mere scavengers; however, detailed research revealed that they are very efficient predators. Information on what spotted hyaenas actually prefer to prey on and what they avoid is lacking, as well as the factors that influence prey selection. Data from 14 published and one unpublished study from six countries throughout the distribution of the spotted hyaena were used to determine which prey species were preferred and which were avoided using Jacobs' index. The mean of these values for each species was used as the dependent variable in multiple regression, with prey abundance and prey body mass as predictive variables. In stark contrast to the rest of Africa's large predator guild, spotted hyaenas do not preferentially prey on any species. Also surprisingly, only buffalo, giraffe and plains zebra are significantly avoided. Spotted hyaena most prefer prey within a body mass range of 56–182 kg, with a mode of 102 kg. The dietary niche breadth of the spotted hyaena is similar to that of the lion Panthera leo , and the two species have a 58.6% actual prey species overlap and a 68.8% preferred prey species overlap. These results highlight the flexible and unselective nature of spotted hyaena predation and are probably a reason for the species' success throughout its range, despite a large degree of dietary overlap with lions.     

Evidence for prey selection by spotted hyaena in the Eastern Cape,South Africa

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.     

Feeding and social behaviour of the striped hyaena (Hyaena vulgaris Desmarest)*

Observations are presented on food, foraging behaviour, ranging and some social behaviour patterns of the striped hyaena; these observations are compared with similar ones on spotted hyaenas. Striped hyaenas are more omnivorous than spotted ones, scavenging a great deal and eating insects, fruits and small vertebrate prey. However, the diets of the two species have several food species in common, and there is competition in the relatively small area where the geographical ranges and habitats of the two hyaenas overlap. Striped hyaenas are solitary, and they show several behavioural adaptations to this. The relationship between the hyaenas' social organization and resource exploitation is discussed.     

Limiting factors of Striped Hyaena,Hyaena hyaena,distribution and densities across climatic and geographical gradients (Mammalia: Carnivora)

Few previous studies on the factors that affect Striped Hyaena (Hyaena hyaena Linnaeus, 1758) occurrence and densities were done on geographically unrelated populations using different methodologies. In Israel, hyaenas occur throughout the country's steep climatic and geographical gradients, presenting a unique opportunity to study densities and habitat use across adjacent ecosystems using a unified methodology and test previous conceptions regarding the species’ habitat selection. We collected hyaena abundance-absence data using 1440 camera traps placed at 80 sites (2012– 2016). Site location ranged from hyper-arid deserts to dense Mediterranean shrubland. We assessed the effect of climate, habitat structure, elevation, geomorphological attributes (proxy for den availability), and anthropogenic development (proximity to settlements and agriculture) on hyaena densities using N-mixture models. Hyaena densities were negatively affected by anthropogenic development, and were limited by den availability. Hyaena densities did not follow a climatic or geographic gradient. Densities were highest at hyper-arid deserts and Mediterranean coastal shrublands. Despite the former conception that hyaenas prefer semi-arid open habitats and avoid extreme deserts and dense vegetation, we show that hyaenas use and even thrive in these habitats when geomorphological conditions are suitable and resources are available.     

Using Landscape and Bioclimatic Features to Predict the Distribution of Lions,Leopards and Spotted Hyaenas in Tanzania's Ruaha Landscape

Tanzania''s Ruaha landscape is an international priority area for large carnivores, supporting over 10% of the world''s lions and important populations of leopards and spotted hyaenas. However, lack of ecological data on large carnivore distribution and habitat use hinders the development of effective carnivore conservation strategies in this critical landscape. Therefore, the study aimed to (i) identify the most significant ecogeographical variables influencing the potential distribution of lions, leopards and spotted hyaenas across the Ruaha landscape; (ii) identify zones with highest suitability for harbouring those species; and (iii) use species distribution modelling algorithms (SDMs) to define important areas for conservation of large carnivores. Habitat suitability was calculated based on environmental features from georeferenced presence-only carnivore location data. Potential distribution of large carnivores appeared to be strongly influenced by water availability; highly suitable areas were situated close to rivers and experienced above average annual precipitation. Net primary productivity and tree cover also exerted some influence on habitat suitability. All three species showed relatively narrow niche breadth and low tolerance to changes in habitat characteristics. From 21,050 km² assessed, 8.1% (1,702 km²) emerged as highly suitable for all three large carnivores collectively. Of that area, 95.4% (1,624 km²) was located within 30 km of the Park-village border, raising concerns about human-carnivore conflict. This was of particular concern for spotted hyaenas, as they were located significantly closer to the Park boundary than lions and leopards. This study provides the first map of potential carnivore distribution across the globally important Ruaha landscape, and demonstrates that SDMs can be effective for understanding large carnivore habitat requirements in poorly sampled areas. This approach could have relevance for many other important wildlife areas that only have limited, haphazard presence-only data, but which urgently require strategic conservation planning.     

Carnivores from the Early Pleistocene hominid-bearing Trinchera Dolina 6 (Sierra de Atapuerca, Spain).

The carnivores from Trinchera Dolina level 6 (TD6) in Sierra de Atapuerca include Ursus sp., Crocuta crocuta, Mustela palerminea, Lynx sp., Canis mosbachensis and Vulpes praeglacialis. Approximately 80 human remains belonging to Homo antecessor were found in the Aurora Stratum (AS): located in the upper part of the TD6 level (TD6, T36-43). The carnivores from AS are taxonomically similar to the rest of the TD6 level, differing only in the occurrence of Lynx sp. The spotted hyaena (Crocuta crocuta) inhabited the Atapuerca Sierra during the Early Pleistocene and up to the Middle Pleistocene, after which it is absent. According to palaeomagnetic, U/Th and ESR results, the fossils from TD6 are dated to the Early Pleistocene, which is the earliest certain occurrence in Europe of Crocuta crocuta. It is associated with Mimomys savini, the arrival of which in Europe can be correlated with the beginning of the Early Biharian. Crocuta crocuta may have arrived during the late Early Pleistocene ("Bavelian complex"), and subsequently invaded the rest of the continent during the Middle and Late Pleistocene. Atapuerca TD6-AS represents the most ancient deposits in Europe where Homo and spotted hyaenas coexisted and, they probably competed ecologically. The carnivores from TD6 are biochronologically consistent with the end of the Early Pleistocene or early Cromerian (corresponding with the Biharian biochron). The timespan could correspond with oxygen isotope stages 19, 20 or 21.     

Spotted hyaena Crocuta crocuta feeding ecology and selectivity of large herbivorous prey in the Namib desert

We have investigated the relationship between spotted hyaenas in the south Namib Desert and large herbivorous prey and have summarized an updated overview of predator‐prey relationships in this resource‐limited arid environment. Over the 52‐month study, we recorded the densities (#/km⁻², ±SE) of the four local large herbivorous prey species: gemsbok (1.229, ±0.50), springbok (1.352, ±0.48), ostrich (0.648, ±0.23), and greater kudu (0.343, ±0.00). A fecal analysis was performed on 146 collected spotted hyaena scats, and prey items were identified and hairs cross‐follicle analyzed to the species level. Spotted hyaena diet at the study area remained opportunistic with 240 identified prey items representing eight differing prey species being recorded, ranging from ostrich eggs to large ungulates. The Ivlev''s Electivity Index was used to determine which large herbivorous prey was most selected for. Although gemsbok had a higher representation of prey items in the sampled scats, all sampled large herbivorous prey species scored below 0 and are thus generally avoided in relation to their availability in the environment. If any prey preferences are expressed by spotted hyaena in the Namib, it can be presumed to be a nonsampled prey species. We therefore promote further detailed investigations into all other prey species present, and seasonal variations of prey densities and scat sampling, within the study environment.     

Diet of striped hyaena in northern Kenya

A study of striped hyaena ( Hyaena hyaena (Linnaeus, 1758)) in Lothagam, northern Kenya was conducted to understand the interactions of H. hyaena with the local Turkana people and their livestock. Data were collected from skeletal parts, fresh scats, ecological survey, and from interviews at different homesteads. Analysis of skeletal remains was broadly divided into three categories: species, skeletal part and bone damage. Insects, birds, fish, crocodile, seeds, leaves and fifteen species of mammalian prey were identified. The high proportion of livestock, dog and human remains in the bone accumulations indicate a significant dependence on the lifestyles of the local Turkana people. The study provides evidence to suggest that striped hyaenas predate on small livestock and demonstrate an opportunistic behaviour, which enables them to survive as the largest carnivore in this marginal environment. A greater abundance of bones are associated with striped hyaena dens than with those of the spotted hyaena ( Crocuta crocuta (Erxleben, 1777)). Bone modification by the striped hyaena differs from spotted hyaena bone modification. Bone breakage patterns can be attributed to the relative specific gravity, degree of epiphyseal closure and nutritive fat content of bones.     

SPOTTED HYAENA WHOOPS: FREQUENT INCIDENCE OF VOCAL INSTABILITIES IN A MAMMALIAN LOUD CALL

ABSTRACT

Spotted hyaena Crocuta crocuta whoops are loud calls normally produced in a sequence termed a bout. Whoops are produced by hyaenas irrespective of age or sex to display identity and convey information about the location of the caller. The majority (91%, n=460) of whoops produced by spotted hyaenas, from two geographically separate populations in southern Africa and one population in eastern Africa, showed pronounced nonlinear phenomena, predominantly subharmonics. Whoops produced by males and females had a similarly high probability of sub- harmonics, and 91.5% of the 78 bouts examined contained calls with subharmonics. These results provide evidence that nonlinear vocal phenomena are a common feature of hyaena whoops. The presence of subharmonics in whoops may be enhanced by vocal tract resonances when the fundamental frequency and first formant in the calls are close or coincide. Vocal membranes may also play a role. The high incidence of subharmonics in whoops may enhance individual recognition by adding structural complexity to calls. As 33 of 34 individually known spotted hyaenas examined in this study produced whoops containing subharmonics, it is unlikely that the production of subharmonics is confined to calls from individuals of a particular social status, sex, size, or level of developmental asymmetry, as proposed for nonlinear phenomena in the calls of other mammalian species, although variation in structural features of subharmonics may convey information about these characteristics.     

Coprolites as a source of information on the genome and diet of the cave hyena

We performed high-throughput sequencing of DNA from fossilized faeces to evaluate this material as a source of information on the genome and diet of Pleistocene carnivores. We analysed coprolites derived from the extinct cave hyena (Crocuta crocuta spelaea), and sequenced 90 million DNA fragments from two specimens. The DNA reads enabled a reconstruction of the cave hyena mitochondrial genome with up to a 158-fold coverage. This genome, and those sequenced from extant spotted (Crocuta crocuta) and striped (Hyaena hyaena) hyena specimens, allows for the establishment of a robust phylogeny that supports a close relationship between the cave and the spotted hyena. We also demonstrate that high-throughput sequencing yields data for cave hyena multi-copy and single-copy nuclear genes, and that about 50 per cent of the coprolite DNA can be ascribed to this species. Analysing the data for additional species to indicate the cave hyena diet, we retrieved abundant sequences for the red deer (Cervus elaphus), and characterized its mitochondrial genome with up to a 3.8-fold coverage. In conclusion, we have demonstrated the presence of abundant ancient DNA in the coprolites surveyed. Shotgun sequencing of this material yielded a wealth of DNA sequences for a Pleistocene carnivore and allowed unbiased identification of diet.     

Feeding ecology and its influence on social organization in Brown hyenas (Hyaena brunnea, Thunberg) of the Central Kalahari Desert

(1) Observations are presented on the diet, feeding habits, hunting and foraging behaviour of Brown hyaenas of the Central Kalahari. (2) The remains of kills left by other predators are the single most important food item in the Brown hyaena's diet. The diet also consists of small scavenged items, small prey such as rodents which the hyaena itself kills, and wild fruits. (3) Brown hyaenas hunt and scavenge small items solitarily, but congregate for communal scavenging of the large kills left by other predators. (4) Individual hyaenas are not territorial and there is great overlap in home ranges. They use common pathways and frequently meet to socialize while foraging. (5) Resident adults form a group with a social hierarchy maintained through neck-biting, muzzle-wrestling, chasing, and other social interactions. Subadult hyaenas often leave the group when approximately 22 months old. (6) Brown hyaenas have a complex system of communication including visual displays, social interactions, vocalizations, and extensive pasting. These are described. (7) Since carrion is an important source of food, Brown hyaenas have developed distinct relationships with other predators and these are described. (8) In conclusion, the Brown hyaena exhibits a highly flexible social system, foraging and hunting small items solitarily and congregating for the common utilization of a large carcass. The social organization is therefore influenced by the feeding ecology.     

The diet of Indian foxes in a peri-urban area of eastern India

Urban expansions into natural habitats have forced many carnivores to adapt to the modified areas; however, our understanding of their ecology in such areas is limited. The Indian fox (Vulpes bengalensis) is a small carnivore species that can adapt and exploit urban areas due to its generalist nature, but there is a paucity of data on Indian foxes in these human-modified areas. To understand the resource requirement of Indian foxes in urban habitats, we conducted a diet analysis by examining 119 scats of Indian foxes in a peri-urban habitat of Baripada town, Odisha, eastern India. To determine the diet composition of Indian foxes, we analyzed scat contents by their frequency occurrence (%FO), the relative frequency of occurrence (%RF) and mean percent volume (%Vm). Indian foxes consumed a wide range of food categories, including arthropods, rodents, reptiles, birds, plant parts and anthropogenic resources. In %Vm, rodents showed the highest volume, followed by polythene, Spondias pinnata fruit and Coleoptera. We found anthropogenic foods in considerable quantities, including polythene and paper. The diet of Indian foxes indicates that the species has opportunistic and generalist feeding habits consisting primarily of easily accessible foods. Furthermore, management efforts should focus on the proper management of waste and control of anthropogenic food sources in urban habitats.     

The ecology of large carnivores in the highlands of northern Ethiopia

The degradation and fragmentation of the northern Ethiopian highlands has resulted in frequent encounters of large carnivores with humans and their livestock. We interviewed 500 randomly selected households to estimate economic impact of livestock predation by spotted hyaena (Crocuta crocuta), leopard (Panthera pardus) and jackal (Canis aureus aureus) in the highlands of northern Ethiopia. The annual mean economic loss per household was approximately U.S.$ 20.2, about 7% of the average annual income of households in the area. Households surveyed reported losses of a total of 3122 livestock to hyaena, leopard and jackal predation over the past 5 years. This loss equated to a total financial loss of U.S.$ 50,381 . Livestock predation incidents of spotted hyaena, leopard and jackal demonstrated that spotted hyaena had a preference for dog, donkey, goat and sheep; leopard for goat, dog and sheep; and jackal for goat and sheep. Livestock predation of spotted hyaena and leopard were mainly during the night. We conclude that assessing depredation problems is important to develop actions for management of either livestock practices or wildlife conservation.     

Boots on the ground: in defense of low-tech,inexpensive, and robust survey methods for Africa’s under-funded protected areas

Protected area managers need reliable information to detect spatial and temporal trends of the species they intend to protect. This information is crucial for population monitoring, understanding ecological processes, and evaluating the effectiveness of management and conservation policies. In under-funded protected areas, managers often prioritize ungulates and carnivores for monitoring given their socio-economic value and sensitivity to human disturbance. Aircraft-based surveys are typically utilized for monitoring ungulates because they can cover large areas regardless of the terrain, but such work is expensive and subject to bias. Recently, unmanned aerial vehicles have shown great promise for ungulate monitoring, but these technologies are not yet widely available and are subject to many of the same analytical challenges associated with traditional aircraft-based surveys. Here, we explore use of inexpensive and robust distance sampling methods in Kafue National Park (KNP) (22,400 km²), carried out by government-employed game scouts. Ground-based surveys spanning 101, 5-km transects resulted in 369 ungulate group detections from 20 species. Using generalized linear models and distance sampling, we determined the environmental and anthropogenic variables influencing ungulate species richness, density, and distribution. Species richness was positively associated with permanent water and percent cover of closed woodland vegetation. Distance to permanent water had the strongest overall effect on ungulate densities, but the magnitude and direction of this effect varied by species. This ground-based approach provided a more cost-effective, unbiased, and repeatable method than aerial surveys in KNP, and could be widely implemented by local personnel across under-funded protected areas in Africa.