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Crocuta

Crocuta is a genus in the family Hyaenidae, comprising the spotted hyena (Crocuta crocuta) as its sole extant species and several extinct species known from the fossil record spanning the Pliocene to Pleistocene epochs, with the genus distributed across Africa, Europe, and Asia.^[1]^[2] The genus originated around 5 million years ago, with fossil evidence indicating a historical range far beyond the current sub-Saharan African distribution of the living species, and morphological adaptations evolving from solitary scavenging to cooperative hunting behaviors.^[1] The spotted hyena, the largest living hyena, measures 95–150 cm in body length, stands 75–85 cm at the shoulder, and weighs 45–90 kg, with females typically larger and more aggressive than males.^[3] It features a sandy to gray coat heavily spotted with black, a sloping back, short hind legs, and exceptionally strong jaws with bone-crushing dentition adapted for scavenging and hunting.^[3] Native to open habitats such as savannas, grasslands, and semi-deserts in sub-Saharan Africa—from Senegal to South Africa—C. crocuta avoids dense forests but can inhabit elevations up to 4,000 m.^[3] Socially, spotted hyenas live in matriarchal clans of 6–100 individuals, where females dominate males and inheritance follows female lines; males disperse from natal clans at maturity to join others.^[3] They are highly vocal, producing a characteristic "whoop" call and a laughter-like sound during conflicts, and exhibit complex behaviors including cooperative hunting of prey like wildebeest and zebras, which forms up to 70% of their diet, supplemented by scavenging.^[3] Reproduction is aseasonal and polygynous, with a 110-day gestation yielding 1–4 cubs (twins common); females possess a unique pseudo-penis formed from fused labia, through which they urinate, mate, and give birth.^[3] Cubs are born precocial but remain dependent in communal dens for up to 18 months.^[3] Ecologically, C. crocuta serves as an apex predator and scavenger, competing with lions and influencing ecosystem dynamics through carcass disposal and nutrient cycling, though it faces human persecution due to livestock depredation.^[3] The species is listed as Least Concern on the IUCN Red List, with a decreasing population trend estimated at 27,000–47,000 individuals, primarily threatened by habitat loss, snares, and retaliatory killings outside protected areas.^[4] Fossil species of Crocuta, such as C. spelaea (the European cave hyena) and up to seven others, highlight the genus's former diversity and adaptation to Ice Age environments before regional extinctions.^[2]

Etymology and taxonomy

Etymology

The genus name Crocuta derives from the Latin crocuta, which in turn is borrowed from the Ancient Greek krokóttas (κροκόττας), referring to a mythical beast described in classical texts as a ferocious dog-wolf hybrid inhabiting Ethiopia or India.^[5] This creature, first attested in the writings of Ctesias (5th–4th century BCE), was said to imitate human voices to lure and devour prey, combining traits like the speed of a horse, the strength of an ox, and the savagery of a lion.^[5] Pliny the Elder further popularized the term in his Naturalis Historia (c. 77 CE), using variants such as crocotas, corocottas, and leucrocottas to describe a similar mythical animal with magical properties, including the ability to dig up gold and speak like humans.^[5] The name krokóttas itself traces etymologically to the Sanskrit koṭṭhâraka (a type of jackal), possibly altered through phonetic shifts in ancient traveler accounts from India to Greece.^[5] These ancient depictions likely stemmed from distorted observations of spotted hyenas (Crocuta crocuta), whose vocalizations and scavenging habits evoked supernatural lore.^[5] The spotted hyena was first described as Canis crocuta by Johann Christian Polycarp Erxleben in his 1777 work Systema Regni Animalis, drawing on earlier descriptions by Thomas Pennant (1771).^[6] The genus Crocuta was established by Johann Jakob Kaup in 1828 to distinguish the spotted hyena from other hyena genera, such as Hyaena (derived from Greek hyaina, meaning "female pig," alluding to the striped and brown hyenas' hog-like features), reflecting early recognition of distinct morphological and behavioral differences within the family Hyaenidae.^[6]^[5]

Taxonomic classification

Crocuta is a genus within the family Hyaenidae, which belongs to the order Carnivora and suborder Feliformia. It is classified in the subfamily Hyaeninae, comprising the bone-cracking hyenas, alongside genera such as Hyaena and Parahyaena.^[7] Phylogenetic analyses based on nuclear and mitochondrial DNA sequences indicate that Crocuta forms a monophyletic clade sister to the group containing Hyaena and Parahyaena, while the aardwolf genus Proteles represents the most basal lineage within Hyaenidae. This positioning reflects the family's evolutionary divergence within Feliformia, closer to felids than canids.^[1] The genus Crocuta is currently monotypic among extant forms, with C. crocuta (the spotted hyena) as its sole living species; no subspecies are recognized due to extensive genetic and morphological variation across populations.^[1] Historically, the spotted hyena was initially described as Canis crocuta by Erxleben in 1777, but morphological distinctions—such as robust build and dentition—led to its separation from the genus Hyaena and the establishment of Crocuta by Kaup in 1828.^[7]^[6]

Species

Extant species

The genus Crocuta is represented by a single extant species, the spotted hyena (Crocuta crocuta), which is the largest living member of the Hyaenidae family. This species inhabits diverse open habitats across sub-Saharan Africa, from semi-deserts and savannas to lightly wooded areas, with confirmed presence in countries including Tanzania, Kenya, Zambia, South Africa, and Ethiopia.^[4]^[3] Its range is widespread across sub-Saharan Africa, though populations are decreasing due to habitat fragmentation.^[4] The spotted hyena possesses a distinctive powerful build, with a massive head, muscular neck and shoulders, and a slightly sloping back that gives it a bear-like appearance. It features a short, dense coat that varies from light brown, beige, or sandy to yellowish-gray, marked by irregular dark spots on the sides, back, and legs; these spots are more prominent in younger individuals and may fade with age. Key identifiers include rounded, short ears and a dog-like muzzle, contributing to its robust skull adapted for bone-crushing. Adults measure 95–165 cm in head-body length and 70–92 cm at the shoulder, with a tail of 25–35 cm.^[8]^[3] Body mass shows sexual dimorphism, with females averaging 49–90 kg and males 40–55 kg, making females the larger sex. Regional variations in size and pelage coloration occur across its range, with individuals in East Africa tending to be larger and those in arid northern areas smaller and paler. Although contemporary taxonomy considers C. crocuta monotypic, earlier classifications proposed various subspecies reflecting observed differences in morphology and coat pattern.^[3]^[8]^[9]

Extinct species

The genus Crocuta includes at least seven extinct species known primarily from fossil records spanning the Pliocene to the Late Pleistocene, distinct from the extant C. crocuta.^[10] One prominent extinct species is Crocuta spelaea, commonly referred to as the cave hyena, which inhabited Eurasia during the Late Pleistocene, approximately 125,000 to 11,700 years ago. Fossils of this species have been extensively recovered from European cave deposits, such as those in Germany, France, and the Czech Republic, where they are often associated with den sites containing bone accumulations from prey like mammoths and reindeer. Morphologically, C. spelaea exhibited a larger body size than modern spotted hyenas, with estimates reaching up to 100 kg based on postcranial measurements, alongside more robust dentition adapted for bone-cracking.^[2]^[11] Another Late Pleistocene species, Crocuta ultima, is recorded from Eurasia, particularly Asia, dating to around 50,000 to 10,000 years ago. Remains of C. ultima have been found in sites such as caves in the Russian Far East and southern Siberia, indicating a distribution across northern latitudes during the Last Glacial Maximum. This species featured larger cheek teeth relative to skull size, suggesting enhanced bone-processing capabilities, though its postcranial skeleton was comparatively smaller than that of C. spelaea, potentially reflecting adaptations to colder environments.^[10]^[12] The earlier Crocuta dietrichi represents a Plio-Pleistocene species from Africa and possibly early Eurasian dispersals, with fossils dated to approximately 3.85–3.63 million years ago in the Pliocene transitioning into the Pleistocene. Key discovery sites include localities in Namibia (e.g., Meob) and Kenya, where mandibular and dental fragments indicate a smaller overall form compared to later Crocuta species, serving as a transitional morphology in the genus's evolutionary history. Its robust dentition, similar to later congeners, points to early specialization in hypercarnivory and scavenging.^[13]

Physical characteristics

Morphology

The genus Crocuta is characterized by a robust body plan adapted for a scavenging and hunting lifestyle, featuring a sloping back resulting from elongated forelegs and shorter hind legs, which elevate the shoulders above the hindquarters. This structure, combined with a massive neck and large head, supports powerful locomotion and prey handling, with body lengths ranging from 95 to 150 cm and shoulder heights of 75 to 85 cm in the extant species.^[3]^[14] The skull of Crocuta is notably massive and specialized for durophagy, with a prominent sagittal crest that provides extensive attachment area for the temporalis muscles, enabling strong bite forces. The dentition includes carnassial teeth (the upper fourth premolar and lower first molar) forming a shearing mechanism, alongside robust premolars designed for bone-crushing, allowing the animal to access marrow from large bones. The dental formula is 3/3, 1/1, 4/3, 1/1, totaling 34 teeth, with the jaws exhibiting the highest strength relative to body size among extant mammals.^[14]^[15] Sensory adaptations in Crocuta emphasize olfaction and vision suited to nocturnal and crepuscular activity. The species possesses an acute sense of smell, facilitated by a large nasal surface area and specialized olfactory behaviors, enabling detection of carrion or prey over long distances. Forward-facing eyes provide binocular vision and enhanced depth perception, aiding in precise hunting strikes.^[16]^[17] The coat of Crocuta consists of coarse, woolly fur with a base color of sandy yellow to gray, overlaid by irregular dark brown or black spots that are most prominent on the neck, shoulders, back, flanks, and legs; these spots are darkest in juveniles and may fade in older adults. Cubs are born with a uniform grayish coat that molts into the spotted pattern by several months of age. A short, erectile mane of longer hairs runs along the neck and back, which can be raised during displays.^[3]^[14] Extinct species within the genus, such as C. spelaea, showed morphological variations including shorter distal limb segments and more robust metapodials compared to the extant form.^[2]

Adaptations

The spotted hyena (Crocuta crocuta), the sole extant species in the genus Crocuta, exhibits remarkable anatomical and physiological adaptations that enhance its survival in competitive savanna ecosystems. These include specialized dentition and cranial features for processing tough food resources, unique reproductive anatomy in females, and traits supporting prolonged physical exertion. One of the most distinctive adaptations is the bone-crushing capability of the jaws, facilitated by a robust skull, powerful masseter muscles, and specialized carnassial teeth with reinforced enamel. This structure allows spotted hyenas to generate bite forces estimated at up to 4,500 N, the highest relative to body size among extant mammals, enabling them to fracture large bones such as those of wildebeest and access nutrient-rich marrow that competitors cannot.^[18] Females possess a highly masculinized external genitalia, characterized by an elongated clitoris resembling a pseudo-penis, which measures up to 17 cm in length and serves multiple functions including urination, mating reception, and parturition. This structure, unique among mammals, results from elevated prenatal androgen exposure and lacks a urogenital sinus typical in other species, requiring dilation during birth to accommodate cubs.^[19]^[20] For endurance during extended pursuits, spotted hyenas have a proportionately large heart, which supports sustained aerobic activity over distances up to 4 km at speeds of 50-60 km/h. Their gait features a flexible spine and overextendable tarsal joints, promoting an energy-efficient "rocking-horse" lope that minimizes fatigue. Thermoregulation is achieved primarily through panting, which facilitates evaporative cooling in hot environments, supplemented by wallowing in water to dissipate heat post-exertion or after large meals.^[21]^[22]^[23]

Distribution and habitat

Current range

The spotted hyena (Crocuta crocuta) is currently distributed across sub-Saharan Africa, with a range extending from Senegal in the west through central and eastern regions to South Africa in the south, encompassing countries including Angola, Botswana, Cameroon, Ethiopia, Kenya, Namibia, Tanzania, Uganda, Zambia, and Zimbabwe. This distribution covers approximately 6.4 million km², though it is patchy in West Africa and more continuous in eastern and southern parts of the continent. The species is notably absent from dense equatorial rainforests, such as the Congo Basin, and extreme desert environments like the central Sahara, where unsuitable conditions limit its presence. Preferred habitats include open savannas, grasslands, semi-deserts, and lightly wooded areas up to elevations of 4,100 meters, where the hyena's scavenging and hunting behaviors are facilitated by visibility and access to prey. These environments often feature seasonal water sources, and the species frequently occurs near human settlements, adapting to anthropogenic landscapes while avoiding heavily forested or arid extremes. Logging activities have occasionally enabled expansion into previously inaccessible woodland edges.^[24] As of the latest assessments, the global population is estimated at 27,000 to 47,000 individuals, with stable or increasing numbers in protected areas like the Serengeti-Mara ecosystem (Tanzania and Kenya) and Kruger National Park (South Africa), which host the largest known clans. Outside these reserves, populations face declines due to habitat fragmentation and conflict, but the overall range remains widespread enough to classify the species as Least Concern.

Historical distribution

The genus Crocuta originated in Africa during the Pliocene, with fossil evidence of early species such as C. dietrichi dating to approximately 3.63–3.85 million years ago in eastern Africa.^[25] Earlier forms remained primarily confined to Africa throughout the Early Pleistocene, as indicated by fossils from sites like Olduvai Gorge in Tanzania.^[26] A major dispersal event occurred around 2.52 million years ago (95% CI: 2.21–2.83 Ma), when ancestral Crocuta populations migrated from Africa into Eurasia, likely via the Sinai-Levant corridor as part of broader faunal exchanges during the Early Pleistocene.^[25] The earliest Eurasian fossils, attributed to C. honanensis, appear in the Longdan Basin of China approximately 2 million years ago, marking the initial expansion into Asia.^[25] By the Middle Pleistocene, around 1 million years ago, Crocuta had reached Europe, with remains of C. crocuta documented at sites like Atapuerca in Spain, facilitating a wide distribution across the continent.^[25] During the Pleistocene, the genus occupied extensive ranges in Eurasia, including the cave hyena (C. crocuta spelaea) in Europe from the Iberian Peninsula to Siberia, and Asian forms like C. crocuta ultima in China and Southeast Asia.^[27] The retraction of Crocuta from Eurasia began in the Late Pleistocene, with populations in central and northern Europe declining around 40,000 years ago, followed by extirpation in western and southern Europe by approximately 31,000 years ago, based on radiocarbon-dated remains.^[28] By about 11,000 years ago, the genus had vanished entirely from Eurasia, coinciding with the end of the Pleistocene and part of the broader Quaternary megafaunal extinction event.^[28] This extinction was likely driven by a combination of climatic deterioration, including increased cold intolerance and reduced prey availability due to declining grassland productivity, as well as heightened competition from humans and other predators like lions and cave bears.^[25]^[28] In Asia, some populations persisted longer, with evidence from Chinese sites indicating survival into the Early Holocene, though ultimately succumbing to similar pressures.^[27]

Biology and ecology

Spotted hyenas (Crocuta crocuta), the sole extant species in the genus Crocuta, live in stable, matriarchal clans typically comprising 6 to 100 individuals, with sizes varying based on resource availability and territory quality.^[29]^[30] These clans are characterized by strict linear dominance hierarchies, where adult females occupy the top ranks and dominate all males, regardless of the males' age or size.^[31]^[30] Rank is inherited matrilineally, with female offspring assuming positions immediately below their mothers, fostering nepotism and coalitionary support among close kin.^[29]^[31] Communication within clans is multifaceted, relying on vocal, olfactory, and visual signals to maintain cohesion, defend territories, and navigate hierarchies. The distinctive "whoop" call, a long-distance vocalization audible up to 5 km, conveys individual identity, sex, and clan affiliation, enabling coordination during territorial disputes or group assemblies.^[31]^[30] Scent marking, primarily via secretions from anal glands pasted onto grass stalks or rubbed on objects, reinforces territorial boundaries and advertises social status, with higher-ranking individuals marking more frequently.^[29]^[30] Females assume leadership roles in clan decision-making and group activities, such as initiating hunts and resolving conflicts, due to their dominant status, which grants them priority access to resources and mates.^[29]^[31] In contrast, males are born into the natal clan but typically emigrate as subadults to join unrelated clans, where they enter at the bottom of the hierarchy and must queue behind other immigrants for potential mating opportunities and status improvement.^[29]^[30] This philopatry in females and dispersal in males promotes genetic diversity while preserving the core matrilineal structure of the clan.^[31]

Diet and foraging

Spotted hyenas (Crocuta crocuta) exhibit an omnivorous diet, with approximately 95% consisting of animal matter obtained through hunting and scavenging, primarily medium to large ungulates such as wildebeest, zebra, and buffalo, including bones, hides, and other tough tissues that most carnivores cannot digest.^[32]^[33] The remaining 5% comprises plant matter, such as grass and occasional fruits, which is incidentally ingested or consumed during periods of food scarcity and appears in fecal samples.^[32] This dietary flexibility allows spotted hyenas to exploit diverse resources across varying habitats, from savannas to semi-arid regions, where ungulate migrations provide seasonal abundance.^[34] Foraging strategies combine active hunting and opportunistic scavenging, with hyenas capable of killing 60–95% of their food intake directly in natural settings, depending on prey availability and location.^[33] Cooperative group hunting enhances success rates, particularly for large prey like wildebeest; small groups of 2–4 individuals achieve up to 31% capture rates for calves by coordinating chases over distances of 500–2,500 meters, targeting vulnerable rumps and loins.^[23] Kleptoparasitism plays a role, as lions frequently steal kills from hyena groups, though hyenas occasionally scavenge abandoned lion kills or engage in limited theft themselves.^[32] A single hyena can consume up to 18 kg of meat in one feeding bout, with clans devouring an entire 100 kg wildebeest carcass in under 15 minutes.^[32] Their digestive system supports this scavenging lifestyle through high efficiency, with a short gut adapted for rapid processing of large volumes of low-quality food, achieving 98–99% nutrient extraction from ingested material.^[35] The stomach produces highly acidic secretions with a pH as low as 1.5, enabling the breakdown of bones, hides, and pathogen-laden carrion while killing bacteria and parasites that could otherwise cause infection.^[36] This combination of morphological adaptations and microbial interactions in the gut minimizes waste, excreting primarily indigestible hair and hooves, and allows hyenas to thrive on diets that would be toxic to other predators.^[35]

Reproduction

Reproduction in the spotted hyena (Crocuta crocuta) is characterized by polyandry, in which receptive females mate with multiple males over short estrus periods, promoting sperm competition and influencing male mating strategies such as harassment and mate guarding.^[37] This polyandrous system occurs within female-dominated clans, where high-ranking females secure preferred mates.^[37] Copulation is uniquely complicated by the female's pseudopenis—an enlarged, erectile clitoris homologous to the male penis—which must be retracted or navigated during intromission, often leading to prolonged and awkward mating attempts.^[38]^[39] Gestation lasts approximately 110 days, after which females give birth to litters of 1–4 cubs, typically 1–2 (twins common).^[40]^[39] Cubs are precocial at birth, emerging fully furred with eyes open and milk teeth already erupted, enabling immediate aggression toward siblings.^[41] However, early cub mortality is high, reaching up to 60% within the first year, primarily due to siblicide, infanticide by unrelated females, starvation, disease, and predation.^[42] Newborn cubs remain largely den-bound for 1-2 months, dependent on maternal lactation while developing foraging skills in the communal den.^[43] They emerge to accompany the clan on hunts around 6 months but continue nursing until 12-18 months.^[44] Sexual maturity is attained at about 3 years, with females typically producing their first litters around this age.^[45] In the wild, spotted hyenas have a lifespan of 12-25 years, though most do not exceed 15-19 years due to environmental pressures.^[30]^[46]

Evolutionary history

Origins and phylogeny

The genus Crocuta originated in Africa during the late Miocene to early Pliocene, approximately 5–7 million years ago, evolving from early hyaenid ancestors within the family Hyaenidae.^[1]^[47] Percrocutids, such as the genus Percrocuta, represent basal hyaenids that appeared in the middle Miocene (around 15–11 million years ago) across Eurasia and Africa, characterized by robust premolars adapted for slicing rather than full bone-crushing. The transition to the Crocuta lineage involved adaptations toward enhanced durophagy, marking a shift from earlier hyaenid forms that relied more on hypercarnivory.^[48] Phylogenetically, Crocuta occupies a basal position within the subfamily Hyaeninae, the bone-crushing hyenas, diverging from other extant hyaenid genera such as Hyaena, Parahyaena, and Proteles around 10 million years ago based on fossil-calibrated molecular estimates.^[13] This divergence reflects an early split in the Hyaeninae clade during the late Miocene, with Crocuta forming a monophyletic group that includes both the modern spotted hyena (C. crocuta) and extinct Eurasian forms like the cave hyena.^[49] Genetic analyses of ancient DNA further support this positioning, showing deep mitochondrial haplogroups within Crocuta that trace back to African origins before dispersals to Eurasia around 2.5 million years ago.^[13] Key fossil evidence from the Pliocene of eastern Africa illustrates the early evolution of Crocuta, particularly the development of specialized bone-crushing dentition. Remains attributed to early species, such as Crocuta eturono from the Turkana Basin in Kenya (dated to approximately 3.8 million years ago), exhibit enlarged carnassial teeth and robust lower carnassials indicative of a transition toward durophagous feeding strategies. Similarly, C. dietrichi fossils from Laetoli, Tanzania (3.85–3.63 million years old), display reinforced jaw structures and conical premolars suited for fracturing bones, distinguishing them from more cursorial Miocene hyaenids.^[13] These African specimens underscore Crocuta's endemic development before its later Pleistocene expansion.^[10]

Extinction events

The genus Crocuta experienced significant range contractions and local extinctions outside Africa during the Late Pleistocene, with Eurasian populations of the cave hyena (C. crocuta or closely related forms) disappearing as part of broader megafaunal turnover. Radiocarbon dating of over 100 specimens indicates extirpation in central Europe and Russia around 40,000 calendar years ago, while northwest and southern European records extend to approximately 31,000 years ago, with no verified survival into the Lateglacial period (post-15,000 years ago).^[50] In eastern Asia, particularly China, the youngest directly dated fossils are from about 20,000 years ago, though population modeling suggests effective extinction occurred between 25,000 and 30,000 years ago following a decline that began around 270,000 years ago and intensified around 60,000 years ago.^[51] These timelines align with the approach to and duration of the Last Glacial Maximum (approximately 26,000–19,000 years ago), after which fossil abundance of Crocuta in Eurasia sharply declined to absence, with no records from the Holocene.^[50]^[51] Multiple interacting factors contributed to these Eurasian extinctions, including physiological limitations to extreme cold, which restricted suitable habitats during glacial advances.^[50] Habitat loss resulted from shifting vegetational zones and reduced primary productivity, leading to prey scarcity for large carnivores like cave hyenas.^[52] Competition intensified with sympatric predators such as cave lions (Panthera spelaea), cave bears (Ursus spelaeus), and gray wolves (Canis lupus), whose ranges overlapped and whose extinctions occurred contemporaneously around 31,000 years ago in western Europe.^[50] Human activities, including hunting and habitat alteration by early modern humans arriving in Europe around 45,000 years ago, likely exacerbated these pressures, particularly in southern refugia where climate remained suitable.^[52]^[50] Species distribution models confirm that while northern Eurasia became climatically unsuitable by the Last Glacial Maximum, southern regions stayed within thermal tolerances, underscoring the role of anthropogenic and ecological factors in the final extirpation.^[52] In Africa, C. crocuta persisted through these global climatic shifts, owing to its behavioral and physiological adaptations to open savanna ecosystems, which buffered against the habitat fragmentation and cooling affecting Eurasian populations.^[51] Genetic analyses reveal that African spotted hyenas maintained higher effective population sizes compared to declining Eurasian lineages, supporting ongoing viability in tropical and subtropical environments.^[51]

Conservation and human relations

Conservation status

The spotted hyena (Crocuta crocuta) is classified as Least Concern on the IUCN Red List (as assessed in 2014), with an estimated population of 27,000 to 47,000 individuals, with more than 10,000 mature individuals, across sub-Saharan Africa.^[4] While overall numbers remain stable in protected areas, populations are declining in fragmented habitats outside these zones due to ongoing anthropogenic pressures, with local subpopulations in eastern, central, and western Africa showing particularly sharp reductions.^[4]^[53] Major threats to the species include habitat loss and fragmentation from agricultural expansion and human settlement, which restrict access to prey and increase encounters with people.^[4] Human-wildlife conflict is rampant, particularly in East Africa, where hyenas predate on livestock, leading to retaliatory killings and persecution.^[54] Poaching for bushmeat, medicinal parts, and due to superstitions associating hyenas with witchcraft further exacerbates mortality, especially in rural communities.^[3] Additionally, hyenas are susceptible to diseases transmitted from domestic livestock, such as anthrax, bovine tuberculosis, and canine distemper, often contracted while scavenging contaminated carcasses.^[55]^[56] Conservation efforts focus on habitat protection and conflict mitigation, with the species safeguarded in key reserves such as Serengeti National Park in Tanzania and Kruger National Park in South Africa, where populations are most secure.^[4]^[53] Community-based programs in East Africa, including predator-proof enclosures and awareness campaigns by organizations like the African Wildlife Foundation, aim to reduce retaliatory killings by compensating farmers and promoting coexistence strategies.^[54]^[57] National action plans, such as Kenya's Recovery and Action Plan for Lion and Spotted Hyena, emphasize monitoring, anti-poaching patrols, and habitat connectivity to support long-term viability.

Interactions with humans

In many African cultures, the spotted hyena (Crocuta crocuta) is depicted in folklore as a mythical scavenger whose distinctive vocalizations resemble laughter, often symbolizing trickery or supernatural cunning.^[58] In Ethiopian traditions, hyenas are closely associated with witchcraft, with beliefs that sorcerers or blacksmiths can transform into these animals to rob graves or perform malevolent acts at night.^[59] Similar negative perceptions prevail in East and Southern African societies, where hyenas are viewed as cowardly thieves or embodiments of evil spirits, reinforcing their role as antagonists in oral stories and rituals.^[60] Historically, humans have utilized spotted hyenas for practical purposes, including hunting them for their skins, which were valued in trade for clothing or trophies, particularly in regions like southern Africa.^[61] Body parts such as sexual organs, rectum, and hairs above the eyes have been employed in traditional medicine across eastern Africa and the Horn, purportedly for love potions, potency enhancement, or treating ailments. Additionally, hyenas have played an inadvertent role in sanitation by scavenging human refuse and corpses, including during conflicts and epidemics, such as dragging bodies amid cholera outbreaks in Somalia or civil wars in Ethiopia, thereby aiding in waste disposal on "battlefields" of disease and war.^[59]^[62] Conflicts between spotted hyenas and humans primarily arise from livestock predation, with hyenas responsible for significant economic losses in pastoral communities across East Africa, such as approximately US$9,600 annually in a community-managed forest in southern Ethiopia due to attacks on cattle and goats.^[63] In retaliation, farmers often resort to poisoning bait, which not only kills hyenas but also affects other wildlife, exacerbating conservation threats from these human-wildlife clashes.^[64] Despite these tensions, spotted hyenas contribute positively to ecotourism in protected areas like national parks in Tanzania and Kenya, where sightings attract visitors and generate revenue through safari experiences that highlight their social behaviors and ecological role.^[61]

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Extreme clitoral development in female spotted hyaenas challenges aspects of the accepted framework of sexual differentiation and reproductive function.
[21]
Human–Hyena (Crocuta crocuta) Conflict in the Tarangire ... - MDPI
Mar 6, 2024 · They have heavy forequarters and a strong large heart to facilitate long distance travel and endurance. Spotted hyenas can cover forty ...
[22]
[PDF] The ecology of spotted hyena,Crocuta crocuta, in Majete Wildlife ...
However, circadian clocks are adaptable and animal behaviour is rather a combination of adaptation to environmental stimuli and circadian clocks (Bloch et al.
[23]
[PDF] Hunting Success in the Spotted Hyena: Morphological Adaptations ...
It is the post-cranial skeleton and muscles of spotted hyenas that set them apart as more efficient cursors than any other extant or extinct bone-cracking ...Missing: plan | Show results with:plan
[24]
Spotted Hyena (Crocuta crocuta) Fact Sheet: Distribution & Habitat
Sep 19, 2025 · Current distribution (from Honer et al. 2008 unless otherwise noted). Africa, south of the Sahara; Native countries.
[25]
Hyena paleogenomes reveal a complex evolutionary history of cross ...
Mar 13, 2020 · We find a deep divergence (~2.5 Ma) between African and Eurasian Crocuta populations, suggesting that ancestral Crocuta left Africa around the same time as ...
[26]
Population History of Extant and Extinct Hyenas - Oxford Academic
Cave hyenas, that is, Pleistocene specimens from Eurasia that belong to the genus Crocuta, are usually considered a subspecies, C. crocuta spelaea, of the ...
[27]
The earliest Middle Pleistocene Crocuta crocuta (Erxleben, 1777) at ...
Jul 26, 2012 · The spotted hyena Crocuta crocuta is an important element of the dispersal events from Africa and Asia to Europe occurred at the Early ...
[28]
Palaeoproteomic analysis of Pleistocene cave hyenas from east Asia
Oct 7, 2020 · Combining the genome data with the oldest Crocuta fossil records in Africa (Crocuta dietrichi, 3.63–3.85 Ma), east Asia (Crocuta ...<|control11|><|separator|>
[29]
New radiocarbon evidence on the extirpation of the spotted hyaena ...
Jul 15, 2014 · The extirpation of spotted hyaena, Crocuta crocuta, in northern Eurasia can be seen as part of the late Quaternary megafaunal extinction event.
[30]
Society, demography and genetic structure in the spotted hyena
Aug 31, 2011 · The stable core of any spotted hyena clan is comprised of one to several matrilineal kin groups (Fig. 1), each containing multiple adult females ...
[31]
Social intelligence in the spotted hyena (Crocuta crocuta) - PMC
These animals have a keen olfactory sense, and they engage in frequent scent-marking behaviour. Each clan appears to have a unique scent signature (Hofer et ...Missing: forward- depth
[32]
What the hyena's laugh tells: Sex, age, dominance and individual ...
Mar 30, 2010 · Spotted hyenas have a matrilineal social system similar to that of many old world primates [40]. Within spotted hyena clans, there are separate ...Missing: matriarchal | Show results with:matriarchal
[33]
Spotted Hyena (Crocuta crocuta) Fact Sheet: Diet & Feeding
Sep 19, 2025 · Scavenging · Contribution to diet varies by locality · Remains often left for extended periods · Sources of carrion.
[34]
Adaptability of large carnivores to changing anthropogenic food ...
Apr 4, 2012 · We previously reported that hyenas get most food from scavenging on urban and rural waste; depredation accounted for <20% of food intake in ...
[35]
Spotted hyaena Crocuta crocuta feeding ecology and selectivity of ...
Mar 11, 2021 · We have investigated the relationship between spotted hyaenas in the south Namib Desert and large herbivorous prey and have summarized an updated overview.
[36]
(PDF) You Gotta Be Shitting Me! Fecal matter composition, digestive ...
Mar 3, 2020 · This study's goal was to determine if there was a correlation between C.crocuta (hereafter referred to as Spotted Hyena or simply, Hyena) digestive efficiency ...Missing: pH 1.5
[37]
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[38]
(PDF) Sexual conflicts in spotted hyenas: Male and female mating ...
Aug 7, 2025 · Polyandry and sperm competition appeared to counter effectively pre-copulatory male tactics, such as harassment, monopolization and other ...
[39]
Sexual Mimicry In Hyenas - ResearchGate
Aug 6, 2025 · This hypothesis stems from data in spotted hyenas, where females develop a pseudopenis that mimics a male penis. Pseudopenis size is positively ...Missing: polyandry | Show results with:polyandry
[40]
Giving birth through a penile clitoris: parturition and dystocia in the ...
Aug 7, 2025 · The average litter consists of 1, 2 or rarely 3 cubs, and the length of the gestation period is 110 days approximately (10). Characteristically, ...
[41]
Hormonal correlates of 'masculinization' in female spotted hyaenas ...
... pregnancy in the spotted hyaena (Crocuta crocuta). The gestation period was estimated to be about 110 days. There was a marked progressive rise in all the …
[42]
Sibling rivalry: training effects, emergence of dominance and ...
Jun 20, 2012 · Aggression between littermates begins within hours after birth as cubs are born with open eyes and teeth erupted [28], results in the ...
[43]
(PDF) Early cub mortality in the spotted hyena, Crocuta crocuta
Nov 18, 2015 · Early cub mortality in the spotted hyena, Crocuta crocuta: effects of maternal rank, communal den use, and maternal favoritism. January 2002.
[44]
The fitness of dispersing spotted hyaena sons is influenced by ... - NIH
Females give birth to one or two, very rarely three, offspring per litter56. Cubs are raised at a communal den, and for the first 6 months of their life depend ...
[45]
Post-weaning maternal effects and the evolution of female ... - NIH
Mar 18, 2009 · Our results demonstrate that maternal support can strongly influence offspring survival beyond weaning in the spotted hyena, a species in which ...Missing: gestation | Show results with:gestation
[46]
[PDF] Society, demography and genetic structure in the spotted hyena
Because the lifespan of wild spotted hyenas may exceed 18 years. (Drea & Frank 2003), but the average age at first repro- duction is only 3.5 years (Holekamp ...
[47]
[PDF] Endurance rivalry among male spotted hyenas: what does it mean to ...
Jul 7, 2022 · Here we found clear support for the idea that male spotted hyenas engage in an endurance rivalry wherein they must typically persist in the ...Missing: adaptations | Show results with:adaptations
[48]
Crocuta crocuta - Plazi TreatmentBank
Mar 13, 2022 · The earliest members of the genus Crocuta first appear in the fossil record of Africa in the early Pliocene, dated at roughly 3-7 million years ...Missing: 1829 | Show results with:1829<|control11|><|separator|>
[49]
Percrocutidae - Wikipedia
The first percrocutids are known from the middle Miocene of Europe and western Asia and belonged to the genus Percrocuta. Percrocuta already had large premolars ...
[50]
[PDF] Evolution of bone-cracking adaptations in hyaenids (Mammalia ...
Primitive hyaenids lacked bone-cracking adaptations. A key innovation was the acquisition of zig-zag Hunter-Schreger bands in the Middle Miocene, marking the ...
[51]
Hyena paleogenomes reveal a complex evolutionary history of cross ...
Mar 13, 2020 · Sardella R., Petrucci M., The earliest Middle Pleistocene Crocuta crocuta (Erxleben, 1777) at Casal Selce (Rome, Italy). Quat. Int. 267, 103 ...
[52]
https://doi.org/10.1016/j.quascirev.2010.04.017
[53]
https://ielc.libguides.com/sdzg/factsheets/spottedhyena/population-conservation
[54]
https://www.awf.org/wildlife-conservation/hyena
[55]
Population & Conservation Status - Spotted Hyena ... - LibGuides
Sep 19, 2025 · Tentative estimate: 27,000-47,000 individuals. · Largest known populations in Serengeti (Tanzania, Kenya) and Kruger National Park · Most stable ...
[56]
Hyena | African Wildlife Foundation
Spotted hyenas are the largest of the three. They are fairly large in build and have relatively short torsos with lower hindquarters, and sloping backs. They ...
[57]
The influence of superstitions and emotions on villagers' attitudes ...
Aug 8, 2023 · We conclude that superstitions can potentially negatively affect hyena persistence, but can be reduced by improving the educational level of local people.Missing: bushmeat | Show results with:bushmeat
[58]
Slow recovery from a disease epidemic in the spotted hyena, a ...
Nov 20, 2018 · CDV infection during the epidemic decreased the survival of cubs and subadults, particularly low-ranking ones. The projected hyena population ...
[59]
Hyena scavenging provides public health and economic benefits to ...
Sep 27, 2021 · They determined that hyena scavenging annually prevents five infections of anthrax and bovine tuberculosis in Mekelle residents and 140 ...
[60]
Kavango Zambezi Transfrontier Conservation Area (KAZA)
Predator-proof livestock enclosures: These structures successfully protect livestock from predators, most often lions and spotted hyenas, by preventing ...
[61]
Hyenas have a bad rap—but they're Africa's most successful predator
Jun 14, 2019 · Myth: Hyenas laugh. ... Vocalizations keep hyena societies intact: Their classic whoop serves to recruit more hyenas during a fight with lions, ...<|control11|><|separator|>
[62]
[PDF] Hyenas and Humans in the Horn of Africa - Everything Harar
Hyenas in the Horn of Africa have a long, ambivalent relationship with humans, both symbiotic and conflicting, and are tolerated for sanitation, but also kill ...
[63]
The Magicality of the Hyena: Beliefs and Practices in West and ... - jstor
ing in eastern central Africa, the spotted hyena is a solar animal that brought the sun to warm the cold earth, but in West Africa it generally symbolizes.
[64]
Ecological determinants of spotted hyena Crocuta crocuta ...
Feb 21, 2024 · According to the World Wildlife Fund, populations of endangered species declined by 60% between 1970 and 2014, and 22 to 36% of known mammal ...
[65]
Ethiopia civil war: Hyenas scavenge on corpses as Tigray forces ...
Oct 18, 2022 · Hyenas scavenging on the corpses of villagers, cities and towns hit in air strikes, elderly men and young women conscripted into armies ...
[66]
Population estimation and livestock loss by spotted hyena (Crocuta ...
In addition, they are widely distributed in eastern, southern, and western Africa (Mills and Hofer, 1998). Spotted hyenas have the most complex social structure ...
[67]
Spotting spotted hyena - Africa Geographic
Jan 23, 2024 · ... hyena are often poisoned to reduce cattle predation. This study demonstrates that the protection offered to hyenas – by buffering human ...Missing: livestock | Show results with:livestock