Hyena Species Research Articles

Striped hyena Hyaena hyaena (Linnaeus 1758): feeding ecology based on den prey remains in a pastoralist landscape, southern Kenya

Abstract Very little is known about the range, diet and eco-ethology of the African striped hyena, compared to its sympatric hyena species; the spotted hyena. Here we present the results of bone assemblages collected in eight striped hyena dens located in a pastoralists’ area in and near the Shompole conservancy, southern Kenya. We present the dens taxonomical representation; comparing domestic and wildlife prey and their prey age structure. This is to aid in making inferences on the striped hyena’s prey acquisition, selection and mode of transportation. Ultimately, this is to highlight the species’ interaction with pastoralists and some eco-ethological features of this little known near threatened species. Total wildlife proportions were higher than livestock. However, both wild and domestic ungulates are the major prey of the striped hyena and carnivores constitute a greater proportion of its non-ungulate prey. Striped hyena appears to predate on younger livestock when their dens are located closer to human settlements than those away from human occupied areas. Juveniles and old adults of wild ungulates dominated the striped hyena dens than the prime adults and this was reversed in the case of livestock ungulates.

Review of the global research on Hyaenidae and implications for conservation and management

Abstract Despite the ecological importance of the four extant species of Hyaenidae, and the threats they face globally, there has been no review of the nearly 100 years of published research on hyaenas, nor has there been a synthesis of management‐related literature regarding these species. We reviewed 907 studies on Hyaenidae, summarized broad temporal, geographic and topical trends, and evaluated findings from management‐related research to determine ways forward for hyaena conservation management. Since the first known study in 1939, most have focused on spotted hyaena (Crocuta crocuta; 75% of all studies), yet overall publications for Hyaenidae have increased by 372% in recent decades. Only 44 of the 67 hyaena range states were represented across publications, with nearly half of all studies conducted in Kenya (18%), South Africa (16%) and Tanzania (13%). Twenty‐eight countries had fewer than five studies. Ecology and diet were the most‐studied topic areas. The least‐studied topics were disease and physiology. Studies on human–hyaena interactions were highly variable in topic, with infrastructure impacts and Hyaenidae benefits to people covered the least. All species were reported to have consumed anthropogenic diet items. Mortality data were included within 11% of publications, with 79% of recorded hyaena mortality constituting anthropogenic causes, although there were few targeted studies on the subject. Lastly, 12% of publications involved community engagement in their methods. There is a significant bias among species, topics and range states across Hyaenidae studies, and little data explicitly related to human–hyaena coexistence. Our management‐focused synthesis suggests that research on Hyaenidae could better reflect large carnivore conservation and management inquiry by increasing studies focused on human interactions with Hyaenidae. To address research gaps and inform Hyaenidae management, we recommend increasing applied research outside of protected areas and using interdisciplinary, community‐involved methods to increase foundational knowledge on understudied hyaena species, habitats and locations.

Seasonal variation in prey preference, diet partitioning and niche breadth in a rich large carnivore guild

AbstractLarge 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.

Leopard density and interspecific spatiotemporal interactions in a hyena-dominated landscape.

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 km2, 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.

Evidence of Toxocara Eggs in Pachycrocuta brevirostris (Gervais, 1850) Coprolites from the Pleistocene of Taurida Cave (Crimea).

Coprolites of the hyena Pachycrocuta brevirostris from the Lower Pleistocene (Upper Villafranchian) of Taurida Cave (Crimea) were studied. One of the three hyena coprolites contained helminth eggs. These eggs were assigned to Toxocara sp. based on their size and morphology. Toxocariasis was evidently a very common infestation among extinct hyena species. The find of toxocara in P. brevirostris coprolite from the Taurida Cave is the earliest evidence of roundworm infestation in P. brevirostris.

Collaboration for conservation: Assessing countrywide carnivore occupancy dynamics from sparse data

AbstractAimAssessing the distribution and persistence of species across their range is a crucial component of wildlife conservation. It demands data at adequate spatial scales and over extended periods of time, which may only be obtained through collaborative efforts, and the development of methods that integrate heterogeneous datasets. We aimed to combine existing data on large carnivores to evaluate population dynamics and improve knowledge on their distribution nationwide.LocationBotswana.MethodsBetween 2010 and 2016, we collated data on African wild dog, cheetah, leopard, brown and spotted hyaena and lion gathered with different survey methods by independent researchers across Botswana. We used a multi‐species, multi‐method dynamic occupancy model to analyse factors influencing occupancy, persistence and colonization, while accounting for imperfect detection. Lastly, we used the gained knowledge to predict the probability of occurrence of each species countrywide.ResultsWildlife areas and communal rangelands had similar occupancy probabilities for most species. Large carnivore occupancy was low in commercial farming areas and where livestock density was high, except for brown hyaena. Lion occupancy was negatively associated with human density; lion and spotted hyaena occupancy was high where rainfall was high, while the opposite applied to brown hyaena. Lion and leopard occupancy remained constant countrywide over the study period. African wild dog and cheetah occupancy declined over time in the south and north, respectively, whereas both hyaena species expanded their ranges. Countrywide predictions identified the highest occupancy for leopards and lowest for the two hyaena species.Main ConclusionsWe highlight the necessity of data sharing and propose a generalizable analytical method that addresses the challenges of heterogeneous data common in ecology. Our approach, which enables a comprehensive multi‐species assessment at large spatial and temporal scales, supports the development of data‐driven conservation guidelines and the implementation of evidence‐based management strategies nationally and internationally.

Phylogenetic Analysis of Striped Hyaena (Hyaena hyaena: Hyaenidae, Carnivora) in Hatay Province in Turkey

Striped hyaena (Hyaena hyaena) is one of the species in danger of extinction and categorized globally as “Under Threatened Organism”. From time to time, different tissue samples and carcasses of the striped hyena are reported in different regions of Anatolia. In this study, 571 bp length of Cytochrome C Oxidase Subunit II (COX 2) of mitochondiral DNA from hair, ears, nails and teeth specimens from six striped hyaenas were amplified and sequenced to determined phylogenetic relationships between close and distant species related to hyaena. Tissue samples using in this study were found randomly at different times in Hatay province, Turkey. According to our results, all colected samples located in Hatay region are the members of H. hyaena species. Moreover, this research is the first molecular research using COX2 gene region for phylogenetic analysis in Turkey. Further investigation can be performed on studies that suggest determining phylogenetic status of striped hyaenas.

Co‐occurrence of high densities of brown hyena and spotted hyena in central Tuli, Botswana

AbstractLarge 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 ± 1.9/100 km2 for brown hyena and 14.9 ± 2.2/100 km2 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.

Census and distribution of large carnivores in the Tsavo national parks, a critical east African wildlife corridor

AbstractKenya's Tsavo National Parks are a critical conservation area and the only wilderness corridor through densely populated human‐dominated landscapes stretching from the East African coast to Lake Victoria, separating extensive undeveloped grasslands south of the equator from the undeveloped semiarid bushlands to the north. At nearly 21,000 km2, Tsavo is one of four lion strongholds in East Africa and part of Africa's most important populations of all other large African carnivores. To provide baseline data on the status of large carnivores, a track survey was undertaken in 2013. Lions and both hyaena species were widely distributed across both parks, while cheetahs and wild dogs were more patchily distributed. Spotted hyaenas were the most abundant, with an estimated population of 3,903 ± 514 (95% CI), followed by lion (706 ± 201), striped hyaena (679 ± 144), leopard (452 ± 98), cheetah (154 ± 74) and wild dog (111 ± 92). Tsavo's carnivores suffer from bushmeat poaching, illegal grazing and retaliatory killing, and increasing human numbers on its periphery. Given Tsavo's importance as a stronghold for lions and an ecological corridor, increased investment in effective protection is strongly recommended. Evaluating trends will require complementary techniques in light of intensive monitoring requirements.

Aardwolf Population Diversity and Phylogenetic Positioning Inferred Using Complete Mitochondrial Genomes

The aardwolf (Proteles cristata) is a unique, insectivorous species of the family Hyaenidae. It occupies a disjunct range in eastern and southern Africa and is possibly a remaining member of a historical dog-like hyaena clade. Although both local and global population status and trends are largely unknown, aardwolves are not uncommon in arid grasslands and the IUCN conservation status of Least Concern appears to be justified. However, they are one of the least studied of the four hyaena species, and we have scarce information about the genetic structure of aardwolf populations. Here, we present the first complete aardwolf mitochondrial genomes and provide novel information about aardwolf evolutionary origins and genetic structure within a single population. Through the investigation of complete mitochondrial genomes from five individuals from a single population within South Africa, we find the mitochondrial diversity of this population to be neither particularly high nor low compared to a number of other mammalian species. Moreover, we also provide additional evidence towards the basal position of the aardwolf within Hyaenidae with a divergence time of 13.0 Ma (95% CI 10.1–16.4 Ma) from all other extant hyaena species.

Capture of a Hyena-Specific Retroviral Envelope Gene with Placental Expression Associated in Evolution with the Unique Emergence among Carnivorans of Hemochorial Placentation in Hyaenidae.

Capture of retroviral envelope genes from endogenous retroviruses has played a role in the evolution of mammals, with evidence for the involvement of these genes in the formation of the maternofetal interface of the placenta. It has been shown that the diversity of captured genes is likely to be responsible for the diversity of placental structures, ranging from poorly invasive (epitheliochorial) to highly invasive (hemochorial), with an intermediate state (endotheliochorial) as found in carnivorans. The latter recapitulate part of this evolution, with the hyena being the sole carnivoran with a hemochorial placenta. In this study, we performed RNA sequencing on hyena placental transcripts and searched for endogenous retroviral envelope genes that have been captured specifically in the Hyaenidae clade and are not found in any other carnivoran. We identified an envelope gene that is expressed in the placenta at the level of the maternofetal interface, as evidenced by in situ hybridization/immunohistochemistry. The gene entry is coincidental with the emergence of the Hyaenidae clade 30 million years ago (Mya), being found at the same genomic locus in all 4 extant hyena species. Its coding sequence has further been maintained during all of Hyaenidae evolution. It is not found in any of the 30 other carnivorans-both Felidae and Canidae-that we screened. This envelope protein does not disclose any fusogenic activity in ex vivo assays, at variance with the syncytin-Car1 gene, which is found in all carnivorans, including the hyena, in which it is still present, transcriptionally active in the placenta, and fusogenic. Together, the present results illustrate the permanent renewal of placenta-specific genes by retroviral capture and de facto provide a candidate gene for the endotheliochorial to hemochorial transition of Hyaenidae among carnivorans.IMPORTANCE The placenta is the most diverse organ among mammals, due in part to stochastic capture of retroviral envelope genes. In carnivorans, capture of syncytin-Car1 took place 80 Mya. It is fusogenic, expressed at the syncytialized placental maternofetal interface, and conserved among all carnivorans, consistent with their shared endotheliochorial placenta. Hyenas are a remarkable exception, with a highly invasive hemochorial placenta, as found in humans, where disruption of maternal blood vessels results in maternal blood bathing the syncytial maternofetal interface. In this study, we identified a retroviral envelope gene capture and exaptation that took place about 30 Mya and is coincident with the emergence of the Hyaenidae, being conserved in all extant hyena species. It is expressed at the maternofetal interface in addition to the shared syncytin-Car1 gene. This new env gene, not present in any other carnivoran, is a likely candidate to be responsible for the specific structure of the hyena placenta.

Sex determining region Y (SRY) sequencing and non-invasive molecular sexing in three wild species: brown (Parahyaena brunnea) and spotted (Crocuta crocuta) hyenas and aardvark (Orycteropus afer).

Non-invasive molecular sexing methods are useful in captive breeding programs and field studies; these methods enable sex identification without immobilisation or stressful handling of animals. We developed a method enabling fast and reliable sex identification in three species with limited external sexual dimorphism: the brown and spotted hyenas and the aardvark. We used the sex determining region Y (SRY ) gene as the male-specific sequence and the c-myc gene, highly conserved among mammals, as the control sequence present in both sexes. Primers designed on the basis of the feline and human SRY gene enabled us to amplify and sequence the SRY gene fragment in hyenas and the aardvark. Subsequently, we used specific primers designed on the basis of the newly obtained sequences for sex determination in two brown hyenas, three spotted hyenas and six aardvarks. The sequences of the SRY gene fragments were further studied. Interspecies comparisons revealed high similarity in SRY sequences between both hyena species, as well as their relationships with the Felidae lineage. The aardvark, as the only species of the order Tubulidentata (Afrotheria), showed SRY gene similarities with Sirenia and Primates. Knowledge of phylogenetic relationships can be beneficial in genetic studies focused on species with limited sequence data.

SERUM BIOCHEMISTRY VALUES AND SELECT SEROLOGIC SCREENING OF BROWN HYENAS ( PARAHYAENA BRUNNEA) FROM THE NAMIB DESERT, NAMIBIA.

Blood from 30 free-ranging brown hyenas ( Parahyaena brunnea) was collected for biochemical analysis and select serologic screening in Namibia from 1997 to 2010. Age was found to have an influence on several biochemical parameters that may be related to growth, a developing immune system, and differences in diet. Seasonal differences in diet of coastal brown hyenas also had an overall significant effect on lipemia values, and differences in stress due to varying capture methods could be associated with an increase in glucose and creatinine kinase. Comparisons among hyena species from published data were inconclusive, as some samples may have been derived from captive populations and individuals. Sera were tested for antibodies against 18 pathogens. Antibodies were not detected for most pathogens, but the proportion of sera containing antibodies against canine adenovirus-1 (CAV-1) and canine adenovirus-2 (CAV-2) was 65% and 84%, respectively. There was no effect of sex, age, year of sampling, or contact with domestic dogs, indicating that CAV-1 or CAV-2 may be enzootic. The prevalence of antibodies to canine distemper virus (CDV) was 43%, and older brown hyenas were 6.9 times more likely to have been exposed to CDV, adjusting for year of sampling and degree of estimated contact with domestic dogs, suggesting epizootic outbreaks. This study is the first to present biochemical reference intervals for wild brown hyenas and provides an indication of disease exposure in this species.

Extended and Continuous Decline in Effective Population Size Results in Low Genomic Diversity in the World's Rarest Hyena Species, the Brown Hyena.

Hyenas (family Hyaenidae), as the sister group to cats (family Felidae), represent a deeply diverging branch within the cat-like carnivores (Feliformia). With an estimated population size of <10,000 individuals worldwide, the brown hyena (Parahyaena brunnea) represents the rarest of the four extant hyena species and has been listed as Near Threatened by the IUCN. Here, we report a high-coverage genome from a captive bred brown hyena and both mitochondrial and low-coverage nuclear genomes of 14 wild-caught brown hyena individuals from across southern Africa. We find that brown hyena harbor extremely low genetic diversity on both the mitochondrial and nuclear level, most likely resulting from a continuous and ongoing decline in effective population size that started ∼1 Ma and dramatically accelerated towards the end of the Pleistocene. Despite the strikingly low genetic diversity, we find no evidence of inbreeding within the captive bred individual and reveal phylogeographic structure, suggesting the existence of several potential subpopulations within the species.

Comparative taphonomical studies of sympatric hyenids (crocuta crocuta and hyaena hyeana) bone assemblages, insights from modern dens in Kenya

Fifty modern dens were investigated to compare bone accumulation patterns of the two hyaena species in two areas in Kenya (Samburu & Shompole). The spotted hyaena bone accumulations were dominated by size class III ungulates while striped hyaenas’ were by size class I & II. Skeletal distribution suggests different accumulation patterns between the two predators, related to their feeding behaviour (hunting versus scavenging). This neo-taphonomy analysis brings additional referential data for the two African hyenids that may be useful for interpretation of bone assemblages and modifications in modern ecology/conservation and would be also applicable to zooarchaeological and paleontological studies.

Çizgili Sırtlanların (Hyaena hyaena) Hatay ve Şanlıurfa‘daki Durumu

Bu çalışmada, Türkiye'de yaklaşık 600 km2'lik alanda 18 ay süresince uygulanan yerli halkla yüz yüze görüşmeler, anket ve foto kapanların yerleştirilmesini içeren bir araştırma gerçekleştirilmiştir. Sahada yapılan anketler ve foto kapan çalışmaları, çizgili sırtlanların Hatay ile Suriye sınırı civarında ve Şanlıurfa‘da güçlü şekilde yaşadıkları konusunda belirgin kanıtlarla sonuçlandı. Ancak, bu bölgede çizgili sırtlanların durumu hakkında kapsamlı bir çalışma yapılmamıştır. Elde edilen sonuçlarımız çizgili sırtlanın sıklıkla çorak arazide ve Hatay ilinde tavuk çiftliği etrafında görüldüğünü göstermiştir. Alternatif olarak, Şanlıurfa İli'nde küçük gruplar halinde yaşıyorlar. Çizgili sırtlanlara uygun birkaç mağara, bu mağaralarda ve sonrasında gözlemlenen ayak izleri, dışkılar ve diğer hayvan kalıntıları ile tespit edilmiştir. Çizgili sırtlanlara uygun birkaç mağara ve sonrasında yapılan gözlemlerde ayak izleri, dışkılar ve diğer hayvan kalıntıları tespit edilmiştir. Saç, kulak ve karkas dokularından izole edilen Cytb mitokondrial DNA'yı kullanarak, çizgili sırtlanların moleküler karakterizasyonunu Türkiye'de ilk defa gerçekleştirdik. Türkiye'den 10 farklı çizgili sırtlan örneğinden Cytb DNA sekanslarının dizilerinin birbirlerinin özdeşi oldukları bulunmuştur. Tarih öncesi olanlar da dahil olmak üzere daha önce rapor edilen Cytb dizilerin karşılaştırılması sonucu Ctyb geninin Hyaena hyaena türleri arasında oldukça muhafaza edildiğini göstermiştir. Arazi çalışması sırasında, yoğun tarım alanları, yerleşimler ve taş ocakları nedeniyle incelenen bölgelerde habitat tahribatının ve parçalanmanın yüksek düzeyde olduğu gözlemlenmiştir. Hatay ve Şanlıurfa illerinde, yasadışı avlanma, çobanlar tarafından sıklıkla mağaranın kullanımı, yaygın kirpi avlanması, zeytinyağı üretim tesislerinden gelen su kirliliği ve karayolu kazaları çizgili sırtlan nüfusunu olumsuz etkilemektedir. Çizgili sırtlanların yaşam alanını korumak için kapsamlı bir çalışma ve koruma planının uygulanması gerektiği sonucuna varılmıştır.

The evolution of intelligence in mammalian carnivores.

Although intelligence should theoretically evolve to help animals solve specific types of problems posed by the environment, it is unclear which environmental challenges favour enhanced cognition, or how general intelligence evolves along with domain-specific cognitive abilities. The social intelligence hypothesis posits that big brains and great intelligence have evolved to cope with the labile behaviour of group mates. We have exploited the remarkable convergence in social complexity between cercopithecine primates and spotted hyaenas to test predictions of the social intelligence hypothesis in regard to both cognition and brain size. Behavioural data indicate that there has been considerable convergence between primates and hyaenas with respect to their social cognitive abilities. Moreover, compared with other hyaena species, spotted hyaenas have larger brains and expanded frontal cortex, as predicted by the social intelligence hypothesis. However, broader comparative study suggests that domain-general intelligence in carnivores probably did not evolve in response to selection pressures imposed specifically in the social domain. The cognitive buffer hypothesis, which suggests that general intelligence evolves to help animals cope with novel or changing environments, appears to offer a more robust explanation for general intelligence in carnivores than any hypothesis invoking selection pressures imposed strictly by sociality or foraging demands.

Spotted, striped or brown? Taphonomic studies at dens of extant hyaenas in eastern and southern Africa

We compare taphonomically relevant aspects of 6 bone accumulations produced by extant spotted hyaenas Crocuta crocuta, striped hyaenas Hyaena hyaena, and brown hyaenas Parahyaena brunnea) from eastern and southern Africa (Republic of Djibouti and Namibia) and published accounts. We highlight similarities and differences in the bone-cracking/consumption and bone collecting habits of each hyaena species with a view to revealing criteria that might be used to distinguish these species. Four parameters are described: i) the prey taken and preferences, ii) the skeletal part distribution, iii) the long bone fragmentation, iv) the tooth mark diversity and frequency. Information from hyaena ecology and the ungulate remains recovered from each bone accumulation reveals differences (or a lack thereof) related to environment, anthropogenic pressure and ethology. Spotted hyaena hunting strategy and clan size suggest a low degree of inter-group competition, but high levels of intra-group on-carcass competition; fewer osteological remains are brought back to dens. In contrast, striped and brown hyaenas are primarily scavengers (removing carcass remnants) and produce significant accumulations of bones at their dens. These differences are likely to be related mainly to inter-specific competition and serve to reduce interaction with other carnivores and to limit kleptoparasitism; the composition of accumulations at dens is determined by food availability and foraging behaviour to a greater or lesser degree. Modern analogues are essential baseline tools for elucidating characteristics that might distinguish each species. The development of neo-taphonomic models is the first step towards comparison and greater understanding of the palaeo-ethology of extinct Pleistocene hyaenids.

Prevalence and diversity of Babesia, Hepatozoon, Ehrlichia, and Bartonella in wild and domestic carnivores from Zambia, Africa

A molecular survey was conducted for several hemoparasites of domestic dogs and three species of wild carnivores from two sites in Zambia. Three Babesia spp. were detected including Babesia felis and Babesia leo in lions (Panthera leo) and a Babesia sp. (similar to Babesia lengau) in spotted hyenas (Crocuta crocuta) and a single lion. All wild dogs (Lycaon pictus) and domestic dogs were negative for Babesia. High prevalences for Hepatozoon were noted in all three wild carnivores (38-61%) and in domestic dogs (13%). Significantly higher prevalences were noted in hyenas and wild dogs compared with domestic dogs and lions. All carnivores were PCR negative for Ehrlichia canis, Ehrlichia ewingii, and Bartonella spp. Overall, high prevalences and diversity of Babesia and Hepatozoon were noted in wild carnivores from Zambia. This study is the first molecular characterization of Babesia from any hyena species and is the first report of a Babesia sp. closely related to B. lengau, a parasite previously only reported from cheetahs (Acinonyx jubatus), in lions and hyenas. Although usually benign in wild carnivores, these hemoparasites can be pathogenic under certain circumstances. Importantly, data on vectors for these parasites are lacking, so studies are needed to identify vectors as well as determine transmission routes, infection dynamics, and host specificity of these hemoparasites in wildlife in Africa and also the risk of transmission between domestic animals and wildlife.

Pleistocene Chinese cave hyenas and the recent Eurasian history of the spotted hyena, Crocuta crocuta

The living hyena species (spotted, brown, striped and aardwolf) are remnants of a formerly diverse group of more than 80 fossil species, which peaked in diversity in the Late Miocene (about 7-8 Ma). The fossil history indicates an African origin, and morphological and ancient DNA data have confirmed that living spotted hyenas (Crocuta crocuta) of Africa were closely related to extinct Late Pleistocene cave hyenas from Europe and Asia. The current model used to explain the origins of Eurasian cave hyena populations invokes multiple migrations out of Africa between 3.5-0.35 Ma. We used mitochondrial DNA sequences from radiocarbon-dated Chinese Pleistocene hyena specimens to examine the origin of Asian populations, and temporally calibrate the evolutionary history of spotted hyenas. Our results support a far more recent evolutionary timescale (430-163 kya) and suggest that extinct and living spotted hyena populations originated from a widespread Eurasian population in the Late Pleistocene, which was only subsequently restricted to Africa. We developed statistical tests of the contrasting population models and their fit to the fossil record. Coalescent simulations and Bayes Factor analysis support the new radiocarbon-calibrated timescale and Eurasian origins model. The new Eurasian biogeographic scenario proposed for the hyena emphasizes the role of the vast steppe grasslands of Eurasia in contrast to models only involving Africa. The new methodology for combining genetic and geological data to test contrasting models of population history will be useful for a wide range of taxa where ancient and historic genetic data are available.