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The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the cl ...
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, currently classed as the sole extant member of the genus ''
Crocuta ''Crocuta'' is a genus of hyena containing the largest living member of the family, the spotted hyena ''(Crocuta crocuta)''. Several fossil species are known as well. Taxonomy It is still unclear whether the genus evolved in Africa or Asia, alth ...
'', native to
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
. It is listed as being of
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. ...
by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
on account of its widespread range and large numbers estimated between 27,000 and 47,000 individuals. The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching. The species may have originated in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, and once ranged throughout
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
for at least one million years until the end of the
Late Pleistocene The Late Pleistocene is an unofficial age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a stratigraphic perspective. It is intended to be the fourth division of the Pleistocene Epoch withi ...
. The spotted hyena is the largest known member of the Hyaenidae, and is further physically distinguished from other species by its vaguely
bear Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the No ...
-like build, its rounded ears, its less prominent mane, its spotted pelt, its more dual-purposed dentition, its fewer
nipple The nipple is a raised region of tissue on the surface of the breast from which, in females, milk leaves the breast through the lactiferous ducts to feed an infant. The milk can flow through the nipple passively or it can be ejected by smooth ...
s and the presence of a pseudo-penis in the female. It is the only
placenta The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate mate ...
l mammalian species where females lack an external vaginal opening, having a pseudo-penis instead. The spotted hyena is the most social of the Carnivora in that it has the largest group sizes and most complex social behaviours. Its social organisation is unlike that of any other carnivore, bearing closer resemblance to that of
cercopithecine The Cercopithecinae are a subfamily of the Old World monkeys, which comprises roughly 71 species, including the baboons, the macaques, and the vervet monkeys. Most cercopithecine monkeys are limited to sub-Saharan Africa, although the macaques ra ...
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter includin ...
s ( baboons and
macaque The macaques () constitute a genus (''Macaca'') of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and (in one instance) Gibraltar. Macaques are principall ...
s) with respect to group size, hierarchical structure, and frequency of social interaction among both kin and unrelated group-mates. However, the social system of the spotted hyena is openly competitive rather than cooperative, with access to kills, mating opportunities and the time of dispersal for males depending on the ability to dominate other
clan A clan is a group of people united by actual or perceived kinship and descent. Even if lineage details are unknown, clans may claim descent from founding member or apical ancestor. Clans, in indigenous societies, tend to be endogamous, mea ...
-members. Females provide only for their own cubs rather than assist each other, and males display no paternal care. Spotted hyena society is
matriarchal Matriarchy is a social system in which women hold the primary power positions in roles of authority. In a broader sense it can also extend to moral authority, social privilege and control of property. While those definitions apply in general En ...
; females are larger than males, and dominate them. The spotted hyena is a highly successful animal, being the most common large carnivore in Africa. Its success is due in part to its adaptability and
opportunism Opportunism is the practice of taking advantage of circumstances – with little regard for principles or with what the consequences are for others. Opportunist actions are expedient actions guided primarily by self-interested motives. The term ...
; it is primarily a hunter but may also scavenge, with the capacity to eat and digest skin, bone and other animal waste. In functional terms, the spotted hyena makes the most efficient use of animal matter of all African carnivores. The spotted hyena displays greater plasticity in its hunting and foraging behaviour than other African carnivores; it hunts alone, in small parties of 2–5 individuals or in large groups. During a hunt, spotted hyenas often run through ungulate herds in order to select an individual to attack. Once selected, their prey is chased over a long distance, often several kilometres, at speeds of up to 60 km/h. The spotted hyena has a long history of interaction with
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
ity; depictions of the species exist from the Upper Paleolithic period, with carvings and paintings from the
Lascaux Lascaux ( , ; french: Grotte de Lascaux , "Lascaux Cave") is a network of caves near the village of Montignac, in the department of Dordogne in southwestern France. Over 600 parietal wall paintings cover the interior walls and ceilings of t ...
and Chauvet Caves. The species has a largely negative reputation in both
Western culture Leonardo da Vinci's ''Vitruvian Man''. Based on the correlations of ideal Body proportions">human proportions with geometry described by the ancient Roman architect Vitruvius in Book III of his treatise ''De architectura''. image:Plato Pio-Cle ...
and African folklore. In the former, the species is mostly regarded as ugly and cowardly, while in the latter, it is viewed as greedy, gluttonous, stupid, and foolish, yet powerful and potentially dangerous. The majority of Western perceptions on the species can be found in the writings of
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
and
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
, though in relatively unjudgmental form. Explicit, negative judgments occur in the ''
Physiologus The ''Physiologus'' () is a didactic Christian text written or compiled in Greek by an unknown author, in Alexandria; its composition has been traditionally dated to the 2nd century AD by readers who saw parallels with writings of Clement of A ...
'', where the animal is depicted as a
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have ...
and grave-robber.Glickman, Stephen (1995)
"The Spotted Hyena from Aristotle to the Lion King: Reputation is Everything In the Company of Animals"
''Social Research'', Volume 62
The IUCN's hyena specialist group identifies the spotted hyena's negative reputation as detrimental to the species' continued survival, both in captivity and the wild.


Etymology and naming

The spotted hyena's scientific name ''Crocuta'', was once widely thought to be derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
loanword A loanword (also loan word or loan-word) is a word at least partly assimilated from one language (the donor language) into another language. This is in contrast to cognates, which are words in two or more languages that are similar because ...
, which translates as 'saffron-coloured one', in reference to the animal's fur colour. This was proven to be incorrect, as the correct spelling of the loanword would have been ''Crocāta'', and the word was never used in that sense by Graeco-Roman sources. ''Crocuta'' actually comes from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
word (), which is derived from the
Sanskrit Sanskrit (; attributively , ; nominally , , ) is a classical language belonging to the Indo-Aryan languages, Indo-Aryan branch of the Indo-European languages. It arose in South Asia after its predecessor languages had Trans-cultural diffusion ...
, which in turn originates from (both of which were originally meant to signify the golden jackal). The earliest recorded mention of Κροκόττας is from
Strabo Strabo''Strabo'' (meaning "squinty", as in strabismus) was a term employed by the Romans for anyone whose eyes were distorted or deformed. The father of Pompey was called " Pompeius Strabo". A native of Sicily so clear-sighted that he could s ...
's ''
Geographica The ''Geographica'' (Ancient Greek: Γεωγραφικά ''Geōgraphiká''), or ''Geography'', is an encyclopedia of geographical knowledge, consisting of 17 'books', written in Greek and attributed to Strabo, an educated citizen of the Roman ...
'', where the animal is described as a mix of wolf and dog native to Ethiopia. From
Classical antiquity Classical antiquity (also the classical era, classical period or classical age) is the period of cultural history between the 8th century BC and the 5th century AD centred on the Mediterranean Sea, comprising the interlocking civilizations of ...
until the
Renaissance The Renaissance ( , ) , from , with the same meanings. is a period in European history marking the transition from the Middle Ages to modernity and covering the 15th and 16th centuries, characterized by an effort to revive and surpass ide ...
, the spotted and striped hyena were either assumed to be the same species, or distinguished purely on geographical, rather than physical grounds.
Hiob Ludolf Hiob or Job Ludolf ( la, Iobus Ludolfus or '; 15 June 1624– 8 April 1704), also known as Job Leutholf, was a German orientalist, born at Erfurt. Edward Ullendorff rates Ludolf as having "the most illustrious name in Ethiopic scholarship". ...
, in his , was the first to clearly distinguish the ''Crocuta'' from '' Hyaena'' on account of physical, as well as geographical grounds, though he never had any first hand experience of the species, having gotten his accounts from an Ethiopian intermediary. Confusion still persisted over the exact taxonomic nature of the hyena family in general, with most European travelers in Ethiopia referring to hyenas as "wolves". This partly stems from the
Amharic Amharic ( or ; (Amharic: ), ', ) is an Ethiopian Semitic language, which is a subgrouping within the Semitic branch of the Afroasiatic languages. It is spoken as a first language by the Amharas, and also serves as a lingua franca for all oth ...
word for hyena, (), which is linked to the
Arabic Arabic (, ' ; , ' or ) is a Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C. E.Watson; Walter ...
word () 'wolf'. The first detailed first-hand descriptions of the spotted hyena by Europeans come from Willem Bosman and Peter Kolbe. Bosman, a Dutch tradesman who worked for the
Dutch West India Company The Dutch West India Company ( nl, Geoctrooieerde Westindische Compagnie, ''WIC'' or ''GWC''; ; en, Chartered West India Company) was a chartered company of Dutch merchants as well as foreign investors. Among its founders was Willem Usselincx ...
at the Gold Coast (modern-day
Ghana Ghana (; tw, Gaana, ee, Gana), officially the Republic of Ghana, is a country in West Africa. It abuts the Gulf of Guinea and the Atlantic Ocean to the south, sharing borders with Ivory Coast in the west, Burkina Faso in the north, and Tog ...
) from 1688 to 1701, wrote of (jackals or woodland dogs) whose physical descriptions match the spotted hyena. Kolben, a
German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ge ...
mathematician and astronomer who worked for the
Dutch East India Company The United East India Company ( nl, Verenigde Oostindische Compagnie, the VOC) was a chartered company established on the 20th March 1602 by the States General of the Netherlands amalgamating existing companies into the first joint-stock ...
in the
Cape of Good Hope The Cape of Good Hope ( af, Kaap die Goeie Hoop ) ;''Kaap'' in isolation: pt, Cabo da Boa Esperança is a rocky headland on the Atlantic coast of the Cape Peninsula in South Africa. A common misconception is that the Cape of Good Hope is ...
from 1705 to 1713, described the spotted hyena in great detail, but referred to it as a "tigerwolf", because the settlers in southern Africa did not know of hyenas, and thus labelled them as "wolves". Bosman and Kolben's descriptions went largely unnoticed until 1771, when the
Welsh Welsh may refer to: Related to Wales * Welsh, referring or related to Wales * Welsh language, a Brittonic Celtic language spoken in Wales * Welsh people People * Welsh (surname) * Sometimes used as a synonym for the ancient Britons (Celtic peopl ...
naturalist
Thomas Pennant Thomas Pennant (14 June OS 172616 December 1798) was a Welsh naturalist, traveller, writer and antiquarian. He was born and lived his whole life at his family estate, Downing Hall near Whitford, Flintshire, in Wales. As a naturalist he had ...
, in his ''Synopsis of Quadrupeds'', used the descriptions, as well as his personal experience with a captive specimen, as a basis for consistently differentiating the spotted hyena from the striped. The description given by Pennant was precise enough to be included by Johann Erxleben in his by simply translating Pennant's text into Latin. ''Crocuta'' was finally recognised as a separate genus from ''Hyaena'' in 1828.


Taxonomy, origins and evolution

Unlike the striped hyena, for which a number of subspecies were proposed in light of its extensive modern range, the spotted hyena is a genuinely variable species, both temporally and spatially. Its range once encompassed almost all of Africa and
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelag ...
, and displayed a large degree of morphological geographic variation, which led to an equally extensive set of specific and subspecific epithets. It was gradually realised that all of this variation could be applied to individual differences in a single subspecies. In 1939, biologist L Harrison Matthews demonstrated through comparisons between a large selection of spotted hyena skulls from Tanzania that all the variation seen in the then recognised subspecies could also be found in a single population, with the only set of characters standing out being pelage (which is subject to a high degree of individual variation) and size (which is subject to
Bergmann's Rule Bergmann's rule is an ecogeographical rule that states that within a broadly distributed taxonomic clade, populations and species of larger size are found in colder environments, while populations and species of smaller size are found in warmer r ...
). When fossils are taken into consideration, the species displayed even greater variation than it does in modern times, and a number of these named fossil species have since been classed as
synonymous A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
with ''Crocuta crocuta'', with firm evidence of there being more than one species within the genus ''Crocuta'' still lacking. Both Björn Kurtén and
Camille Arambourg Camille Arambourg ( February 3, 1885– November 19, 1969) was a French vertebrate paleontologist. He conducted extensive field work in North Africa. In the 1950s he argued against the prevailing model of Neanderthals as brutish and simian. Du ...
promoted an Asiatic origin for the species; Kurtén focussed his arguments on the
Plio-Pleistocene The Plio-Pleistocene is an informally described geological pseudo-period, which begins about 5 million years ago (Mya) and, drawing forward, combines the time ranges of the formally defined Pliocene and Pleistocene epochs—marking from about 5&n ...
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
''Crocuta sivalensis'' from the
Siwaliks The Sivalik Hills, also known as the Shivalik Hills and Churia Hills, are a mountain range of the outer Himalayas that stretches over about from the Indus River eastwards close to the Brahmaputra River, spanning the northern parts of the Ind ...
, a view defended by Arambourg, who nonetheless allowed the possibility of an Indo-Ethiopian origin. This stance was contested by Ficarelli and Torre, who referred to evidence of the spotted hyena's presence from African deposits dating from the early Pleistocene, a similar age to the Asian ''C. sivalensis''. Studies on the phylogeographic distribution of mtDNA
haplotype A haplotype ( haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material ( DNA) which is inherited from two parents. Normally these organisms have their DNA o ...
s indicates three migration events from Africa to Eurasia, though neither the topology of the
phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
or the fossil record exclude the possibility of an Asian origin. The earliest migration of spotted hyenas from Africa to Eurasia began less than 3.5 million years ago, most probably from the area where the first spotted hyena fossils were discovered, reaching
East Asia East Asia is the eastern region of Asia, which is defined in both geographical and ethno-cultural terms. The modern states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. China, North Korea, South Korea ...
and most likely also
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
. The second migration of spotted hyenas occurred less than 1.3–1.5 million years ago and resulted in the first arrival of hyenas in Europe and a separation of African spotted hyenas into a southern and a northern population. The third spotted hyena migration took place 0.36 million years ago, starting from the northern African population and reaching both Europe and Asia. Unlike other African carnivores, with the exception of the
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, ...
, there is no evidence to suggest that spotted hyenas underwent a genetic bottleneck during the Pleistocene. The ancestors of the genus ''Crocuta'' diverged from ''Hyaena'' (the genus of striped and brown hyenas) 10 million years ago. The ancestors of the spotted hyena probably developed social behaviours in response to increased pressure from other predators on carcasses, which forced them to operate in teams. At one point in their
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
, spotted hyenas developed sharp carnassials behind their crushing premolars; this rendered waiting for their prey to die no longer a necessity, as is the case for brown and striped hyenas, and thus they became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory and long chases in a small territory would have caused them to encroach into another clan's land. It has been theorised that female dominance in spotted hyena clans could be an adaptation in order to successfully compete with males on kills, and thus ensure that enough milk is produced for their cubs. Another theory is that it is an adaptation to the length of time it takes for cubs to develop their massive skulls and jaws, thus necessitating greater attention and dominating behaviours from females. Its appearance in Europe and China during the Cromerian period coincided with the decline and eventual extinction of '' Pachycrocuta brevirostris'', the giant short-faced hyena. As there is no evidence of environmental change being responsible, it is likely that the giant short-faced hyena became extinct due to competition with the spotted hyena.


Description


Anatomy

The spotted hyena has a strong and well developed neck and forequarters, but relatively underdeveloped hindquarters. The rump is rounded rather than angular, which prevents attackers coming from behind from getting a firm grip on it. The head is wide and flat with a blunt muzzle and broad rhinarium. In contrast to the striped hyena, the ears of the spotted hyena are rounded rather than pointed. Each foot has four digits, which are webbed and armed with short, stout and blunt claws. The paw-pads are broad and very flat, with the whole undersurface of the foot around them being naked. The tail is relatively short, being long, and resembles a pompom in appearance. Unusually among hyaenids, and mammals in general, the female spotted hyena is considerably larger than the male. Both sexes have a pair of
anal gland Anal may refer to: Related to the anus *Related to the anus of animals: ** Anal fin, in fish anatomy ** Anal vein, in insect anatomy ** Anal scale, in reptile anatomy *Related to the human anus: ** Anal sex, a type of sexual activity involving ...
s which open into the
rectum The rectum is the final straight portion of the large intestine in humans and some other mammals, and the gut in others. The adult human rectum is about long, and begins at the rectosigmoid junction (the end of the sigmoid colon) at the l ...
just inside the anal opening. These glands produce a white, creamy secretion which is pasted onto grass stalks by everting the rectum. The odour of this secretion is very strong, smelling of boiling cheap
soap Soap is a salt of a fatty acid used in a variety of cleansing and lubricating products. In a domestic setting, soaps are surfactants usually used for washing, bathing, and other types of housekeeping. In industrial settings, soaps are us ...
or burning, and can be detected by humans several metres downwind. The spotted hyena has a proportionately large
heart The heart is a muscular Organ (biology), organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as ca ...
, constituting close to 1% of its body weight, thus giving it great endurance in long chases. In contrast, a lion's heart makes up only 0.45–0.57 percent of its body weight. The now extinct Eurasian populations were distinguished from the modern African populations by their shorter distal extremities and longer humerus and
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
. The skull of the spotted hyena differs from that of the striped hyena by its much greater size and narrower
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exception ...
. For its size, the spotted hyena has one of the most powerfully built skulls among the Carnivora. The dentition is more dual purposed than that of other modern hyena species, which are mostly scavengers; the upper and lower third premolars are conical bone-crushers, with a third bone-holding cone jutting from the lower fourth premolar. The spotted hyena also has its carnassials situated behind its bone-crushing premolars, the position of which allows it to crush bone with its premolars without blunting the carnassials. Combined with large jaw muscles and a special vaulting to protect the skull against large forces, these characteristics give the spotted hyena a powerful bite which can exert a pressure of 80  kgf/cm2 (1140
lbf/in² The pound per square inch or, more accurately, pound-force per square inch (symbol: lbf/in2; abbreviation: psi) is a unit of pressure or of stress based on avoirdupois units. It is the pressure resulting from a force of one pound-force applied ...
), which is 40% more force than a leopard can generate. The jaws of the spotted hyena outmatch those of the
brown bear The brown bear (''Ursus arctos'') is a large bear species found across Eurasia and North America. In North America, the populations of brown bears are called grizzly bears, while the subspecies that inhabits the Kodiak Islands of Alaska is k ...
in bone-crushing ability, and free ranging hyenas have been observed to crack open the long bones of
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, '' Giraffa camelopardal ...
s measuring 7 cm in diameter. A spotted hyena is estimated to have a bite force of 565.7 newtons at the canine tip and 985.5 newtons at the carnassial eocene. One individual in a study was found to exert a bite force of 4,500 newtons on the measuring instruments.


Dimensions

The spotted hyena is the largest extant member of the Hyaenidae. Adults measure in body length, and have a shoulder height of . Adult male spotted hyenas in the
Serengeti The Serengeti ( ) ecosystem is a geographical region in Africa, spanning northern Tanzania. The protected area within the region includes approximately of land, including the Serengeti National Park and several game reserves. The Serenget ...
weigh , while females weigh . Spotted hyenas in
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most central point. Its neighbours are t ...
tend to be heavier, with males weighing on average , and females . Exceptionally large weights of and are known. It has been estimated that adult members of the now extinct Eurasian populations weighed .


Fur

Fur colour varies greatly and changes with age. Unlike the fur of the striped and brown hyena, that of the spotted hyena consists of spots rather than stripes and is much shorter, lacking the well defined spinal mane of the former two species. The base colour generally is a pale greyish-brown or yellowish-grey on which an irregular pattern of roundish spots is superimposed on the back and hind quarters. The spots, which are of variable distinction, may be reddish, deep brown or almost blackish. The spots vary in size, even on single individuals, but are commonly in diameter. A less distinct spot pattern is present on the legs and belly but not on the throat and chest. Some research groups (such as Ngorongoro Crater Hyena Project and MSU Hyena Project) often use the spot patterns to help identify individual hyenas). A set of five, pale and barely distinct bands replace the spots on the back and sides of the neck. A broad, medial band is present on the back of the neck, and is lengthened into a forward facing crest. The crest is mostly reddish-brown in colour. The crown and upper part of the face is brownish, save for a white band above both eyes, though the front of the eyes, the area around the rhinarium, the lips and the back portion of the chin are all blackish. The limbs are spotted, though the feet vary in colour, from light brown to blackish. The fur is relatively sparse and consists of two hair types; moderately fine underfur (measuring ) and long, stout bristle hairs (). European Paleolithic rock art depicting the species indicates that the Eurasian populations retained the spots of their modern-day African counterparts.


Female genitalia

The genitalia of the female closely resembles that of the male; the
clitoris The clitoris ( or ) is a female sex organ present in mammals, ostriches and a limited number of other animals. In humans, the visible portion – the glans – is at the front junction of the labia minora (inner lips), above the op ...
is shaped and positioned like a penis, a pseudo-penis, and is capable of
erection An erection (clinically: penile erection or penile tumescence) is a physiological phenomenon in which the penis becomes firm, engorged, and enlarged. Penile erection is the result of a complex interaction of psychological, neural, vascular, a ...
. The female also possesses no external
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
(vaginal opening), as the labia are fused to form a pseudo-
scrotum The scrotum or scrotal sac is an anatomical male reproductive structure located at the base of the penis that consists of a suspended dual-chambered sac of skin and smooth muscle. It is present in most terrestrial male mammals. The scrotum co ...
. The pseudo-penis is traversed to its tip by a central urogenital canal, through which the female urinates, copulates and gives birth. The pseudo-penis can be distinguished from the males' genitalia by its somewhat shorter length, greater thickness, and more rounded glans.Glickman SE, Cunha GR, Drea CM, Conley AJ and Place NJ. (2006).
Mammalian sexual differentiation: lessons from the spotted hyena
'. Trends Endocrinol Metab 17:349–356.
In both males and females, the base of the glans is covered with penile spines. The formation of the pseudo-penis appears largely androgen independent, as the pseudo-penis appears in the female fetus before differentiation of the fetal ovary and adrenal gland. When flaccid, the pseudo-penis is retracted into the abdomen, and only the prepuce is visible. After giving birth, the pseudo-penis is stretched, and loses many of its original aspects; it becomes a slack-walled and reduced prepuce with an enlarged orifice with split lips.


Behaviour


Social behaviour

Spotted hyenas are social animals that live in large communities (referred to as "
clan A clan is a group of people united by actual or perceived kinship and descent. Even if lineage details are unknown, clans may claim descent from founding member or apical ancestor. Clans, in indigenous societies, tend to be endogamous, mea ...
s") which can consist of at most 80 individuals. Group-size varies geographically; in the Serengeti, where prey is migratory, clans are smaller than those in the Ngorongoro Crater, where prey is sedentary. Spotted hyena clans are more compact and unified than
wolf The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
packs, but are not as closely knit as those of African wild dogs. Females usually dominate males, including in cases where low-ranking females generally dominate over high-ranking males, but they will also occasionally co-dominate with a male. There have also been cases in which a clan has been led by a male rather than a female. Cubs take the rank directly below their mothers at birth. So when the matriarch passes away (or, in rare instances, disperses into another clan), their youngest female cub will take over as matriarch. It is typical for females to remain with their natal clan (although there have been rare instances of them dispersing), thus large clans usually contain several matrilines, whereas males typically disperse from their natal clan at the age of 2½ years. Höner et al. say that when a male co-dominates with a female or is otherwise able to lead, this is because the male was born to the matriarch of the clan and has taken the rank directly below his mother. The clan is a fission-fusion society, in which clan-members do not often remain together, but may forage alone or in small groups. High-ranking hyenas maintain their position through aggression directed against lower-ranking clan-members. Spotted hyena
hierarchy A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
is nepotistic; the offspring of dominant females automatically outrank adult females subordinate to their mother. However, rank in spotted hyena cubs is greatly dependent on the presence of the mother; low-ranking adults may act aggressively toward higher-ranking cubs when the mother is absent. Although individual spotted hyenas only care for their own young, and males take no part in raising their young, cubs are able to identify relatives as distantly related as great-aunts. Also, males associate more closely with their own daughters rather than unrelated cubs, and the latter favor their fathers by acting less aggressively toward them. Spotted hyena societies are more complex than those of other carnivorous mammals, and are remarkably similar to those of
cercopithecine The Cercopithecinae are a subfamily of the Old World monkeys, which comprises roughly 71 species, including the baboons, the macaques, and the vervet monkeys. Most cercopithecine monkeys are limited to sub-Saharan Africa, although the macaques ra ...
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter includin ...
s in respect to group size, structure, competition and cooperation. Like cercopithecine primates, spotted hyenas use multiple sensory modalities, recognise individual conspecifics, are conscious that some clan-mates may be more reliable than others, recognise third-party kin and rank relationships among clan-mates, and adaptively use this knowledge during social decision making. Also, like cercopithecine primates, dominance ranks in hyena societies are not correlated with size or aggression, but with ally networks.Drea CM, Frank LG (2003) The social complexity of spotted hyenas. In: ''Animal Social Complexity: Intelligence, Culture, and Individualized Societies'' (eds de Waal FBM, Tyack PL). pp. 121–148, Harvard University Press, Cambridge, Massachusetts. In this latter trait, the spotted hyena further show parallels with primates by acquiring rank through
coalition A coalition is a group formed when two or more people or groups temporarily work together to achieve a common goal. The term is most frequently used to denote a formation of power in political or economical spaces. Formation According to ''A Gui ...
. However, rank reversals and overthrows in spotted hyena clans are very rare. The
social network A social network is a social structure made up of a set of social actors (such as individuals or organizations), sets of dyadic ties, and other social interactions between actors. The social network perspective provides a set of methods fo ...
dynamics of spotted hyenas are determined by multiple factors. Environmental factors include rainfall and prey abundance; individual factors include preference to bond with females and with kin; and topological effects include the tendency to close triads in the network. Female hyenas are more flexible than males in their social bonding preferences. Higher ranking adult spotted hyenas tend to have higher
telomere A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes. Although there are different architectures, telomeres, in a broad sense, are a widespread genetic feature mos ...
length and higher levels of some immune defense proteins in their blood serum. Territory size is highly variable, ranging from less than 40 km2 in the Ngorongoro Crater to over 1,000 km2 in the Kalahari. Home ranges are defended through vocal displays, scent marking and boundary patrols. Clans mark their territories by either pasting or pawing in special latrines located on clan range boundaries. Clan boundaries are usually respected; hyenas chasing prey have been observed to stop dead in their tracks once their prey crosses into another clan's range. Hyenas will however ignore clan boundaries in times of food shortage. Males are more likely to enter another clan's territory than females are, as they are less attached to their natal group and will leave it when in search of a mate. Hyenas travelling in another clan's home range typically exhibit bodily postures associated with fear, particularly when meeting other hyenas. An intruder can be accepted into another clan after a long period of time if it persists in wandering into the clan's territory, dens or kills.


Mating, reproduction, and development

The spotted hyena is a non-seasonal breeder, though a birth peak does occur during the
wet season The wet season (sometimes called the Rainy season) is the time of year when most of a region's average annual rainfall occurs. It is the time of year where the majority of a country's or region's annual precipitation occurs. Generally, the sea ...
. Females are polyestrous, with an estrus period lasting two weeks. Like many feliform species, the spotted hyena is promiscuous, and no enduring pair bonds are formed. Members of both sexes may copulate with several mates over the course of several years. Males will show submissive behaviour when approaching females in heat, even if the male outweighs his partner. Females usually favour younger males born or joined into the clan after they were born. Older females show a similar preference, with the addition of preferring males with whom they have had long and friendly prior relationships. Passive males tend to have greater success in courting females than aggressive ones. Copulation in spotted hyenas is a relatively short affair, lasting 4–12 minutes, and typically only occurs at night with no other hyenas present. The mating process is complicated, as the male's penis enters and exits the female's reproductive tract through her pseudo-penis rather than directly through the vagina, which is blocked by the false scrotum and testes. These unusual traits make mating more laborious for the male than in other mammals, while also ensuring that forced copulation is physically impossible. Once the female retracts her clitoris, the male enters the female by sliding beneath her, an operation facilitated by the penis's upward angle. Once this is accomplished, a typical mammalian mating posture is adopted. Copulation may be repeated multiple times during a period of several hours. Both partners usually lick their genitals for several minutes after mating. The length of the gestation period tends to vary greatly, though 110 days is the average length of time. In the final stages of pregnancy, dominant females provide their developing offspring with higher androgen levels than lower-ranking mothers do. The higher androgen levels the result of high concentrations of ovarian
androstenedione Androstenedione, or 4-androstenedione (abbreviated as A4 or Δ4-dione), also known as androst-4-ene-3,17-dione, is an endogenous weak androgen steroid hormone and intermediate in the biosynthesis of estrone and of testosterone from dehy ...
are thought to be responsible for the extreme
masculinization Virilization or masculinization is the biological development of adult male characteristics in young males or females. Most of the changes of virilization are produced by androgens. Virilization is most commonly used in three medical and biology ...
of female behavior and morphology. This has the effect of rendering the cubs of dominant females more aggressive and sexually active than those of lower ranking hyenas; high ranking male cubs will attempt to mount females earlier than lower ranking males. The average litter consists of two cubs, with three occasionally being reported. Males take no part in the raising of young. Giving birth is difficult for female hyenas, as the females give birth through their narrow clitoris, and spotted hyena cubs are the largest carnivoran young relative to their mothers' weight. During parturition, the clitoris ruptures in order to facilitate the passage of the young, and may take weeks to heal. Cubs are born with soft, brownish black hair, and weigh 1.5 kg on average. Unique among carnivorous mammals, spotted hyenas are also born with their eyes open and with 6–7 mm long canine teeth and 4 mm long incisors. Also, cubs will attack each other shortly after birth. This is particularly apparent in same sexed litters, and can result in the death of the weaker cub. This neonatal siblicide kills an estimated 25% of all hyenas in their first month. Male cubs which survive grow faster and are likelier to achieve reproductive dominance, while female survivors eliminate rivals for dominance in their natal clan. Lactating females can carry of milk in their udders. Spotted hyena milk has the highest
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
and fat content of any terrestrial carnivore. Cubs will nurse from their mother for 12–16 months, though they can process solid food as early as three months. Mothers do not regurgitate food for their young. Females are very protective of their cubs, and will not tolerate other adults, particularly males, approaching them. Spotted hyenas exhibit adult behaviours very early in life; cubs have been observed to ritually sniff each other and mark their living space before the age of one month. Within ten days of birth, they are able to move at considerable speed. Cubs begin to lose the black coat and develop the spotted, lighter coloured pelage of the adults at 2–3 months. They begin to exhibit hunting behaviours at the age of eight months, and will begin fully participating in group hunts after their first year. Spotted hyenas reach sexual maturity at the age of three years. The average lifespan in zoos is 12 years, with a maximum of 25 years.


Denning behaviour

The clan's social life revolves around a communal den. While some clans may use particular den sites for years, others may use several different dens within a year or several den sites simultaneously. Spotted hyena dens can have more than a dozen entrances, and are mostly located on flat ground. The tunnels are usually oval in section, being wider than they are high, and narrow down from an entrance width of ½–1 metre (1.6–7.7 ft) to as small as . In the rocky areas of East Africa and Congo, spotted hyenas use caves as dens, while those in the Serengeti use kopjes as resting areas in daylight hours. Dens have large bare patches around their entrances, where hyenas move or lie down on. Because of their size, adult hyenas are incapable of using the full extent of their burrows, as most tunnels are dug by cubs or smaller animals. The structure of the den, consisting of small underground channels, is likely an effective anti-predator device which protects cubs from predation during the absence of the mother. Spotted hyenas rarely dig their own dens, having been observed for the most part to use the abandoned burrows of warthogs, springhares and jackals. Faeces are usually deposited away from the den, though they urinate wherever they happen to be. Dens are used mostly by several females at once, and it is not uncommon to see up to 20 cubs at a single site. The general form of a spotted hyena den is tunnel-shaped, with a spacious end chamber used for sleeping or breeding. This chamber measures up to in width, the height being rather less. Females generally give birth at the communal den or a private birth den. The latter is primarily used by low status females in order to maintain continual access to their cubs, as well as ensure that they become acquainted with their cubs before transferral to the communal den.


Intelligence

Compared to other hyenas, the spotted hyena shows a greater relative amount of
frontal cortex The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere (in front of the parietal lobe and the temporal lobe). It is parted from the parietal lobe by a groove be ...
which is involved in the mediation of social behavior. Studies strongly suggest
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
in spotted hyena and primate intelligence. A study done by evolutionary anthropologists demonstrated that spotted hyenas outperform
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...
s on cooperative problem-solving tests; captive pairs of spotted hyenas were challenged to tug two ropes in unison to earn a food reward, successfully cooperating and learning the maneuvers quickly without prior training. Experienced hyenas even helped inexperienced clan-mates to solve the problem. In contrast, chimps and other primates often require extensive training, and cooperation between individuals is not always as easy for them.LiveScience Staff (28 September 2009
"Hyenas Surprisingly Good at Cooperative Tasks"
/ref> The intelligence of the spotted hyena was attested to by Dutch colonists in 19th-century South Africa, who noted that hyenas were exceedingly cunning and suspicious, particularly after successfully escaping from traps. Spotted hyenas seem to plan on hunting specific species in advance; hyenas have been observed to engage in activities such as scent marking before setting off to hunt zebras, a behaviour which does not occur when they target other prey species. Also, spotted hyenas have been recorded to utilise deceptive behaviour, including giving alarm calls during feeding when no enemies are present, thus frightening off other hyenas and allowing them to temporarily eat in peace. Similarly, mothers will emit alarm calls when attempting to interrupt attacks on their cubs by other hyenas.


Ecology


Diet

The spotted hyena is the most carnivorous member of the Hyaenidae. Unlike its brown and striped cousins, the spotted hyena is a predator, not a scavenger; this has been shown since the 1960s. One of the earliest studies to demonstrate their hunting abilities was done by Hans Kruuk, an African wildlife ecologist, and he showed through a 7-year study of hyena populations in Africa that spotted hyenas hunt as much as lions, and with later studies this has been shown to be the average in all areas of Africa. However spotted hyenas remain mislabeled as scavengers, often even by ecologists and wildlife documentary channels.
Wildebeest Wildebeest ( , , ), also called gnu ( or ), are antelopes of the genus ''Connochaetes'' and native to Eastern and Southern Africa. They belong to the family Bovidae, which includes true antelopes, cattle, goats, sheep, and other even-toe ...
are the most commonly taken medium-sized ungulate prey item in both Ngorongoro and the Serengeti, with zebra and Thomson's gazelles coming close behind. Cape buffalo are rarely attacked due to differences in habitat preference, though adult bulls have been recorded to be taken on occasion. In Kruger National Park,
blue wildebeest The blue wildebeest (''Connochaetes taurinus''), also called the common wildebeest, white-bearded gnu or brindled gnu, is a large antelope and one of the two species of wildebeest. It is placed in the genus '' Connochaetes'' and family Bovidae, a ...
, cape buffalo,
Burchell's zebra Burchell's zebra (''Equus quagga burchellii'') is a southern subspecies of the plains zebra. It is named after the British explorer and naturalist William John Burchell. Common names include bontequagga, Damaraland zebra, and Zululand zebra (Gray, ...
, greater kudu and impala are the spotted hyena's most important prey, while
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, '' Giraffa camelopardal ...
, impala, wildebeest and zebra are its major food sources in the nearby
Timbavati The Timbavati Private Nature Reserve is located at the border line between Hoedspruit (in Limpopo) and Acornhoek (in Mpumalanga), north of the Sabi Sand Private Game Reserve on the western edge of Kruger National Park. Geographically and polit ...
area. Springbok and kudu are the main prey in
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
's
Etosha National Park Etosha National Park is a national park in northwestern Namibia and one of the largest national parks in Africa. It was proclaimed a game reserve in March 1907 in Ordinance 88 by the Governor of German South West Africa, Friedrich von Lindequist ...
, and springbok in the Namib. In the southern Kalahari,
gemsbok The gemsbok or South African oryx (''Oryx gazella'') is a large antelope in the genus ''Oryx''. It is native to the extremely dry, arid regions of Southern Africa; notably, the Kalahari Desert. Some authorities formerly classified the East Afr ...
, wildebeest and springbok are the principal prey. In Chobe, the spotted hyena's primary prey consists of migratory zebra and resident impala. In
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
's
Masai Mara Maasai Mara, also sometimes spelled Masai Mara and locally known simply as The Mara, is a large national game reserve in Narok, Kenya, contiguous with the Serengeti National Park in Tanzania. It is named in honor of the Maasai people, the ancest ...
, 80% of the spotted hyena's prey consists of
topi ''Damaliscus lunatus jimela'' is a subspecies of topi, and is usually just called a topi. It is a highly social and fast type of antelope found in the savannas, semi-deserts, and floodplains of sub-Saharan Africa. Names The word ''tope'' or ''t ...
and Thomson's gazelle, save for during the four-month period when zebra and wildebeest herds migrate to the area.
Bushbuck The Cape bushbuck (''Tragelaphus sylvaticus'') is a common and a widespread species of antelope in sub-Saharan Africa.Wronski T, Moodley Y. (2009)Bushbuck, harnessed antelope or both? ''Gnusletter'', 28(1):18-19. Bushbuck are found in a wide ran ...
, suni and buffalo are the dominant prey items in the Aberdare Mountains, while Grant's gazelle, gerenuk,
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to domesticate ...
,
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s and
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Adult females are referred to as cows and adult ...
are likely preyed upon in northern Kenya. In west Africa, the spotted hyena is primarily a scavenger who will occasionally attack domestic stock and medium-size antelopes in some areas. In
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the ...
, it is common for spotted hyenas to feed on small antelopes like kob, but may also scavenge on reedbuck, kongoni, buffalo, giraffe, African elephant, topi and roan antelope carcasses. Records indicate that spotted hyenas in
Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northe ...
feed on medium to large-sized ungulates such as
waterbuck The waterbuck (''Kobus ellipsiprymnus'') is a large antelope found widely in sub-Saharan Africa. It is placed in the genus '' Kobus'' of the family Bovidae. It was first described by Irish naturalist William Ogilby in 1833. Its 13 subspecies a ...
and impala. In Tanzania's
Selous Game Reserve The Selous Game Reserve, now renamed Nyerere National Park, is a protected area in southern Tanzania. It covers a total area of and has additional buffer zones. It was designated a UNESCO World Heritage Site in 1982 due to its wildlife diversity ...
, spotted hyenas primarily prey on wildebeest, followed by buffalo, zebra, impala, giraffe, reedbuck and kongoni. In
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The ...
, it is thought that the species primarily preys on birds and reptiles, while in
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most central point. Its neighbours are t ...
it is considered a scavenger.


Enemies and competitors


Lions

Where spotted hyenas and lions occupy the same geographic area, the two species occupy the same ecological niche, and are thus in direct competition with one another. In some cases, the extent of dietary overlap can be as high as 68.8%. Lions typically ignore spotted hyenas, unless they are on a kill or are being harassed by them. There exists a common misconception that hyenas steal kills from lions, but most often it is the other way around, and lions will readily steal the kills of spotted hyenas. In the Ngorongoro Crater, it is common for lions to subsist largely on kills stolen from hyenas. Lions are quick to follow the calls of hyenas feeding, a fact demonstrated by field experiments, during which lions repeatedly approached whenever the tape-recorded calls of hyenas feeding were played. When confronted on a kill by lions, spotted hyenas will either leave or wait patiently at a distance of 30–100 metres until the lions have finished eating. In some cases, spotted hyenas are bold enough to feed alongside lions, and may occasionally force lions off a kill. This mostly occurs during the nighttime, when hyenas are bolder. Spotted hyenas usually prevail against groups of lionesses unaccompanied by males if they outnumber them 4:1. In some instances they were seen to have taken on and routed two pride males while outnumbering them 5:1. The two species may act aggressively toward one another even when there is no food at stake. Lions may charge at hyenas and maul them for no apparent reason; one male lion was filmed killing two hyenas on separate occasions without eating them, and lion predation can account for up to 71% of hyena deaths in
Etosha Etosha National Park is a national park in northwestern Namibia and one of the largest national parks in Africa. It was proclaimed a game reserve in March 1907 in Ordinance 88 by the Governor of German South West Africa, Friedrich von Lindequist. ...
. Spotted hyenas have adapted to this pressure by frequently mobbing lions which enter their territories. Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to the sight of them, but will react fearfully to the scent.


Cheetahs and leopards

Although cheetahs and leopards preferentially prey on smaller animals than those hunted by spotted hyenas, hyenas will steal their kills when the opportunity presents itself. Cheetahs are usually easily intimidated by hyenas, and put up little resistance, while leopards, particularly males, may stand up to hyenas. There are records of some male leopards preying on hyenas. Hyenas are nonetheless dangerous opponents for leopards; there is at least one record of a young adult male leopard dying from a
sepsis Sepsis, formerly known as septicemia (septicaemia in British English) or blood poisoning, is a life-threatening condition that arises when the body's response to infection causes injury to its own tissues and organs. This initial stage is follo ...
infection caused by wounds inflicted by a spotted hyena. There is also a case of two spotted hyenas killing and eating a young male leopard in
Timbavati Game Reserve The Timbavati Private Nature Reserve is located at the border line between Hoedspruit (in Limpopo) and Acornhoek (in Mpumalanga), north of the Sabi Sand Private Game Reserve on the western edge of Kruger National Park. Geographically and polit ...
, apparently in revenge after a young hyena was killed by the leopard.


African wild dogs

Spotted hyenas will follow packs of African wild dogs in order to appropriate their kills. They will typically inspect areas where wild dogs have rested and eat any food remains they find. When approaching wild dogs at a kill, solitary hyenas will approach cautiously and attempt to take off with a piece of meat unnoticed, though they may be mobbed by the dogs in the attempt. When operating in groups, spotted hyenas are more successful in pirating dog kills, though the dog's greater tendency to assist each other puts them at an advantage against spotted hyenas, who rarely work in unison. Cases of dogs scavenging from spotted hyenas are rare. Although wild dog packs can easily repel solitary hyenas, on the whole, the relationship between the two species is a one sided benefit for the hyenas, with wild dog densities being negatively correlated with high hyena populations.


Jackals and wolves

Black-backed and side-striped jackals, and African wolves will feed alongside hyenas, though they will be chased if they approach too closely. Spotted hyenas will sometimes follow jackals and wolves during the gazelle fawning season, as jackals and wolves are effective at tracking and catching young animals. Hyenas do not take to eating wolf flesh readily; four hyenas were reported to take half an hour in eating a golden wolf. Overall, the two animals typically ignore each other when there is no food or young at stake.


Other competitors

Though they readily take to water to catch and store prey, spotted hyenas will avoid crocodile-infested waters, and usually keep a safe distance from
Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed throughout sub-Saharan Africa, occurring mostly in the central, eastern, ...
s. Recent observations shows that African rock pythons can hunt adult spotted hyenas.


Other hyena species

Spotted hyenas dominate other hyena species wherever their ranges overlap. Brown hyenas encounter spotted hyenas in the Kalahari, where the brown outnumber the spotted. The two species typically encounter each other on carcasses, which the larger spotted species usually appropriate. Sometimes, brown hyenas will stand their ground and raise their manes while emitting growls. This usually has the effect of seemingly confusing spotted hyenas, which will act bewildered, though they will occasionally attack and maul their smaller cousins. Similar interactions have been recorded between spotted and striped hyenas in the Serengeti.


Communication


Body language

Spotted hyenas have a complex set of postures in communication. When afraid, the ears are folded flat, and are often combined with baring of the teeth and a flattening of the mane. When attacked by other hyenas or by wild dogs, the hyena lowers its hindquarters. Before and during an assertive attack, the head is held high with the ears cocked, mouth closed, mane erect and the hindquarters high. The tail usually hangs down when neutral, though it will change position according to the situation. When a high tendency to flee an attacker is apparent, the tail is curled below the belly. During an attack, or when excited, the tail is carried forward on the back. An erect tail does not always accompany a hostile encounter, as it has also been observed to occur when a harmless social interaction occurs. Although they do not wag their tails, spotted hyenas will flick their tails when approaching dominant animals or when there is a slight tendency to flee. When approaching a dominant animal, subordinate spotted hyenas will walk on the knees of their forelegs in submission. Greeting ceremonies among clan-members consist of two individuals standing parallel to each other and facing opposite directions. Both individuals raise their hind legs and lick each other's anogenital area. During these greeting ceremonies, the penis or pseudo-penis often becomes erect, in both males and females. Erection is usually a sign of submission, rather than dominance, and is more common in males than in females.


Vocalisations

The spotted hyena has an extensive vocal range, with sounds ranging from whoops, fast whoops, grunts, groans, lows, giggles, yells, growls, soft grunt-laughs, loud grunt-laughs, whines and soft squeals. The loud ' call, along with the maniacal laughter, are among the most recognisable sounds of Africa. Typically, very high-pitched calls indicate fear or submission, while loud, lower-pitched calls express aggression. The pitch of the laugh indicates the hyena's age, while variations in the frequency of notes used when hyenas make noises convey information about the animal's social rank. Dr. Hans Kruuk compiled the following table on spotted hyena calls in 1972;


Diseases and parasites

Spotted hyenas may contract brucellosis, rinderpest and
anaplasmosis Anaplasmosis is a tick-borne disease affecting ruminants, dogs, and horses, and is caused by ''Anaplasma'' bacteria. Anaplasmosis is an infectious but not contagious disease. Anaplasmosis can be transmitted through mechanical and biological vector ...
. They are vulnerable to '' Trypanosoma congolense'', which is contracted by consuming already infected herbivores, rather than through direct infection from tsetse flies. It is known that adult spotted hyenas in the Serengeti have
antibodies An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of ...
against
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, ...
, canine herpes, canine brucellosis,
canine parvovirus Canine parvovirus (also referred to as CPV, CPV2, or parvo) is a contagious virus mainly affecting dogs. CPV is highly contagious and is spread from dog to dog by direct or indirect contact with their feces. Vaccines can prevent this infectio ...
, feline calicivirus,
leptospirosis Leptospirosis is a blood infection caused by the bacteria '' Leptospira''. Signs and symptoms can range from none to mild (headaches, muscle pains, and fevers) to severe ( bleeding in the lungs or meningitis). Weil's disease, the acute, sever ...
, bovine brucellosis, rinderpest and anaplasmosis. During the canine distemper outbreak of 1993–94, molecular studies indicated that the viruses isolated from hyenas and lions were more closely related to each other than to the closest canine distemper virus in dogs. Evidence of canine distemper in spotted hyenas has also been recorded in the Masai Mara. Exposure to rabies does not cause clinical symptoms or affect individual survival or longevity. Analyses of several hyena saliva samples showed that the species is unlikely to be a rabies vector, thus indicating that the species catches the disease from other animals rather than from intraspecifics. The
microfilaria ::''Microfilaria may also refer to an informal "collective group" genus name, proposed by Cobbold in 1882. While a convenient category for newly discovered microfilariae which can not be assigned to a known species because the adults are unknown, ...
of ''
Dipetalonema dracuneuloides Setariidae is a family of nematodes belonging to the order Spirurida Spirurida is an order of spirurian nematodes. Like all nematodes, they have neither a circulatory nor a respiratory system. Some Spirurida, like the genus '' Gongylonema'', ...
'' have been recorded in spotted hyenas in northern Kenya. The species is known to carry at least three
cestode Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, known as tapeworms. Their bodies consist of ...
species of the genus '' Taenia'', none of which are harmful to humans. It also carries
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
n parasites of the genus ''
Hepatozoon ''Hepatozoon'' is a genus of Apicomplexa alveolates which incorporates over 300 species obligate intraerythrocytic parasites. Species have been described from all groups of tetrapod vertebrates, as well as a wide range of haematophagous arthropod ...
'' in the Serengeti, Kenya and South Africa. Spotted hyenas may act as hosts in the life-cycles of various parasites which start life in herbivores; '' Taenia hyaenae'' and ''T. olnogojinae'' occur in hyenas in their adult phase. '' Trichinella spiralis'' are found as cysts in hyena muscles.


Range, habitat and population

The spotted hyena's distribution once ranged in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
from the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, def ...
to the
Urals The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through western ...
, where it remained for at least one million years. Remains have also been found in the
Russian Far East The Russian Far East (russian: Дальний Восток России, r=Dal'niy Vostok Rossii, p=ˈdalʲnʲɪj vɐˈstok rɐˈsʲiɪ) is a region in Northeast Asia. It is the easternmost part of Russia and the Asian continent; and is admin ...
, and it has been theorised that the presence of hyenas there may have delayed the colonisation of North America. The causes of the species' extinction in Eurasia are still largely unknown. In
Western Europe Western Europe is the western region of Europe. The region's countries and territories vary depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the ancient Mediterranean ...
at least, the spotted hyena's extinction coincided with a decline in
grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...
s 12,500 years ago. Europe experienced a massive loss of lowland habitats favoured by spotted hyenas, and a corresponding increase in
mixed woodland Temperate broadleaf and mixed forest is a temperate climate terrestrial habitat type defined by the World Wide Fund for Nature, with broadleaf tree ecoregions, and with conifer and broadleaf tree mixed coniferous forest ecoregions. These fo ...
s. Spotted hyenas, under these circumstances, would have been outcompeted by
wolves The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
and humans which were as much at home in forests as in open lands, and in highlands as in lowlands. Spotted hyena populations began to shrink roughly 20,000 years ago, completely disappearing from Western Europe between 14 and 11,000 years ago, and earlier in some areas.C. Stiner, Mary (2004
Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy
''Revue de Paléobiologie, Genève''
Historically, the spotted hyena was widespread throughout
Sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
. It is present in all habitats save for the most extreme desert conditions, tropical rainforests and the top of alpine mountains. Its current distribution is patchy in many places, especially in West Africa. Populations are concentrated in protected areas and surrounding land. There is a continuous distribution over large areas of
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
,
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
,
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
,
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label= Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kal ...
,
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
and the Transvaal Lowveld areas of
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
. During the 1770s and 1780s the species was still widespread in southern and western South Africa, being recorded on the
Cape Peninsula The Cape Peninsula ( af, Kaapse Skiereiland) is a generally mountainous peninsula that juts out into the Atlantic Ocean at the south-western extremity of the African continent. At the southern end of the peninsula are Cape Point and the Ca ...
and
Cape Flats The Cape Flats ( af, Die Kaapse Vlakte) is an expansive, low-lying, flat area situated to the southeast of the central business district of Cape Town. The Cape Flats is also the name of an administrative region of the City of Cape Town, which li ...
, and near present-day Somerset West, Riviersonderend, Mossel Bay, George,
Joubertina Joubertina is a small town in the Kou-Kamma Local Municipality, Sarah Baartman District of the Eastern Cape province of South Africa. Description Town on the Wabooms River in the Langkloof, some 50 km north-west of Assegaaibos, 70  ...
,
Gamtoos River Gamtoos River or Gamptoos River is a river in the Eastern Cape Province, South Africa. It is formed by the confluence of the Kouga River and the Groot River and is approximately long with a catchment area of . Course The Gamtoos river system ...
,
Jansenville Jansenville is a town in Sarah Baartman District Municipality in the Eastern Cape province of South Africa. Town on the Sundays River, 87 km south of Graaff-Reinet. Laid out on the farm Vergenoegd in 1854, it was proclaimed in 1855 and b ...
,
Cannon Rocks Cannon Rocks is a small seaside town on the Indian Ocean in the Eastern Cape Province, South Africa, between Boknes and the boundary of the Greater Addo Elephant Park. It is situated 135 km east of Port Elizabeth and 180 km west of ...
,
Alice Alice may refer to: * Alice (name), most often a feminine given name, but also used as a surname Literature * Alice (''Alice's Adventures in Wonderland''), a character in books by Lewis Carroll * ''Alice'' series, children's and teen books by ...
,
Onseepkans Onseepkans is a small settlement on the banks of the Orange River in Northern Cape Province, South Africa. It is a border post with Namibia for traffic between Pofadder in South Africa and Keetmanshoop in Namibia. The name either originated fro ...
and
Augrabies Falls The Augrabies Falls is a waterfall on the Orange River, the largest river in South Africa. Since 1966 the waterfall, set in a desolate and rugged milieu, is enclosed by the Augrabies Falls National Park. The falls are around in height. Some s ...
. The species dwells in semi-
desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
s,
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
, open
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the '' plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...
, dense dry woodland, and mountainous forests up to 4,000 m in altitude. It is scarce or absent in
tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equator ...
s and
coast The coast, also known as the coastline or seashore, is defined as the area where land meets the ocean, or as a line that forms the boundary between the land and the coastline. The Earth has around of coastline. Coasts are important zones in n ...
al areas. Its preferred habitats in west Africa include the Guinea and Sudan savannahs, and is absent in the belt of dense coastal forest. In the Namib Desert, it occurs in riverine growth along seasonal rivers, the sub-desertic pro-Namib and the adjoining inland plateau. In ideal habitats, the spotted hyena outnumbers other large carnivores, including other hyena species. However, the striped and brown hyena occur at greater densities than the spotted species in desert and semi-desert regions.
Population densities Population density (in agriculture: standing stock or plant density) is a measurement of population per unit land area. It is mostly applied to humans, but sometimes to other living organisms too. It is a key geographical term.Matt RosenberPo ...
based on systematic censuses vary substantially, from 0.006 to 1.7 individuals per km2.


Relationships with humans


Cultural depictions

In Africa, the spotted hyena is usually portrayed as an abnormal and ambivalent animal, considered to be sly, brutish,
necrophagous Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding ...
and dangerous. It further embodies physical power, excessiveness, ugliness, stupidity, as well as sacredness. Spotted hyenas vary in their folkloric and mythological depictions, depending on the ethnic group from which the tales originate. It is often difficult to know whether or not spotted hyenas are the specific hyena species featured in such stories, particularly in West Africa, as both spotted and striped hyenas are often given the same names. In west African tales, spotted hyenas symbolise immorality, dirty habits, the reversal of normal activities, and other negative traits, and are sometimes depicted as bad
Muslim Muslims ( ar, المسلمون, , ) are people who adhere to Islam, a monotheistic religion belonging to the Abrahamic tradition. They consider the Quran, the foundational religious text of Islam, to be the verbatim word of the God of Abrah ...
s who challenge the local
animism Animism (from Latin: ' meaning ' breath, spirit, life') is the belief that objects, places, and creatures all possess a distinct spiritual essence. Potentially, animism perceives all things— animals, plants, rocks, rivers, weather syst ...
that exists among the Beng in
Côte d’Ivoire Ivory Coast, also known as Côte d'Ivoire, officially the Republic of Côte d'Ivoire, is a country on the southern coast of West Africa. Its capital is Yamoussoukro, in the centre of the country, while its largest city and economic centre is ...
. In East Africa, Tabwa mythology portrays the spotted hyena as a solar animal that first brought the sun to warm the cold earth. Spotted hyenas feature prominently in the rituals of certain African cultures. In the Gelede cult of the
Yoruba people The Yoruba people (, , ) are a West African ethnic group that mainly inhabit parts of Nigeria, Benin, and Togo. The areas of these countries primarily inhabited by Yoruba are often collectively referred to as Yorubaland. The Yoruba constitut ...
of Benin and Southwest Nigeria, a spotted hyena mask is used at dawn to signal the end of the ceremony. As the spotted hyena usually finishes the meals of other carnivores, the animal is associated with the conclusion of all things. Among the Korè cult of the
Bambara people The Bambara ( bm, ߓߡߊߣߊ߲, italics=no, ''Bamana'' or ''Banmana'') are a Mandé ethnic group native to much of West Africa, primarily southern Mali, Guinea, Burkina Faso and Senegal. They have been associated with the historic Bambara Empi ...
in Mali, the belief that spotted hyenas are hermaphrodites appears as an ideal in-between in the ritual domain. The role of the spotted hyena mask in their rituals is often to turn the neophyte into a complete moral being by integrating his male principles with femininity. The Beng people believe that upon finding a freshly killed hyena with its anus inverted, one must plug it back in, for fear of being struck down with perpetual laughter. They also view spotted hyena faeces as contaminating, and will evacuate a village if a hyena relieves itself within village boundaries. In
Harar Harar ( amh, ሐረር; Harari: ሀረር; om, Adare Biyyo; so, Herer; ar, هرر) known historically by the indigenous as Gey (Harari: ጌይ ''Gēy'', ) is a walled city in eastern Ethiopia. It is also known in Arabic as the City of Saint ...
, Ethiopia, spotted hyenas are regularly fed by the city's inhabitants, who believe the hyenas' presence keeps devils at bay, and associate mystical properties such as fortune telling to them. Traditional Western beliefs about the spotted hyena can be traced back to
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
's '' Historia Animalium'', which described the species as a
necrophagous Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding ...
, cowardly and potentially dangerous animal. He further described how the hyena uses retching noises to attract dogs. In ''
On the Generation of Animals The ''Generation of Animals'' (or ''On the Generation of Animals''; Greek: ''Περὶ ζῴων γενέσεως'' (''Peri Zoion Geneseos''); Latin: ''De Generatione Animalium'') is one of the biological works of the Corpus Aristotelicum, the co ...
'', Aristotle criticised the erroneous belief that the spotted hyena is a
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have ...
(which likely originated from the confusion caused by the masculinised genitalia of the female), though his physical descriptions are more consistent with the striped hyena.
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
supported Aristotle's depiction, though he further elaborated that the hyena can imitate human voices. Additionally, he wrote how the hyena was held in high regard among the
Magi Magi (; singular magus ; from Latin '' magus'', cf. fa, مغ ) were priests in Zoroastrianism and the earlier religions of the western Iranians. The earliest known use of the word ''magi'' is in the trilingual inscription written by Darius t ...
, and that hyena body parts could cure different diseases, give protection and stimulate sexual desire in people. Natural historians of the 18th and 19th centuries rejected stories of hermaphroditism in hyenas, and recognised the differences between the spotted and striped hyena. However, they continued to focus on the species' scavenging habits, their potential to rob graves and their perceived cowardice. During the 20th century, Western and African stereotypes of the spotted hyena converged; in both
Ernest Hemingway Ernest Miller Hemingway (July 21, 1899 – July 2, 1961) was an American novelist, short-story writer, and journalist. His economical and understated style—which he termed the iceberg theory—had a strong influence on 20th-century f ...
's ''
Green Hills of Africa ''Green Hills of Africa'' is a 1935 work of nonfiction by American writer Ernest Hemingway. Hemingway's second work of nonfiction, ''Green Hills of Africa'' is an account of a month on safari he and his wife, Pauline Marie Pfeiffer, took in East ...
'' and
Disney The Walt Disney Company, commonly known as Disney (), is an American multinational mass media and entertainment conglomerate headquartered at the Walt Disney Studios complex in Burbank, California. Disney was originally founded on October ...
's ''
The Lion King ''The Lion King'' is a 1994 American animated musical drama film produced by Walt Disney Feature Animation and released by Walt Disney Pictures. The 32nd Disney animated feature film and the fifth produced during the Disney Renaissance ...
'', the traits of gluttony and comical stupidity, common in African depictions of hyenas, are added to the Western perception of hyenas being cowardly and ugly. After the release of ''The Lion King'', hyena biologists protested against the animal's portrayal: one hyena researcher sued Disney studios for
defamation of character Defamation is the act of communicating to a third party false statements about a person, place or thing that results in damage to its reputation. It can be spoken (slander) or written (libel). It constitutes a tort or a crime. The legal defi ...
, and another who had organized the animators' visit to the
University of California The University of California (UC) is a public land-grant research university system in the U.S. state of California. The system is composed of the campuses at Berkeley, Davis, University of California, Irvine, Irvine, University of Califor ...
's Field Station for Behavioural Research, where they would observe and sketch captive hyenas suggested
boycott A boycott is an act of nonviolent, voluntary abstention from a product, person, organization, or country as an expression of protest. It is usually for moral, social, political, or environmental reasons. The purpose of a boycott is to inflict so ...
ing the film.


Livestock predation

When targeting livestock, the spotted hyena primarily preys upon cattle, sheep and goats, though hyenas in the southern parts of
Tigray Region The Tigray Region, officially the Tigray National Regional State, is the northernmost regional state in Ethiopia. The Tigray Region is the homeland of the Tigrayan, Irob, and Kunama people. Its capital and largest city is Mekelle. Tigray ...
of
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
preferentially target
donkey The domestic donkey is a hoofed mammal in the family Equidae, the same family as the horse. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as ...
s. Reports of livestock damage are often not substantiated, and hyenas observed scavenging on a carcass may be mistaken for having killed the animal. The rate at which the species targets livestock may depend on a number of factors, including stock keeping practices, the availability of wild prey and human-associated sources of organic material, such as rubbish. Surplus killing has been recorded in South Africa's eastern Cape Province. Attacks on stock tend to be fewer in areas where livestock is corralled by thorn fences and where domestic dogs are present. One study in northern Kenya revealed that 90% of all cases of livestock predation by hyenas occurred in areas outside the protection of thorn fences.


Attacks on humans and grave desecration

Like most mammalian predators, the spotted hyena is typically shy in the presence of humans, and has the highest
flight distance The flight zone of an animal is the area surrounding an animal that if encroached upon by a potential predator or threat, including humans, will cause alarm and escape behavior. The flight zone is determined by the animal's flight distance, someti ...
(up to 300 metres) among African carnivores. However, this distance is reduced during the night, when hyenas are known to follow people closely. Although spotted hyenas do prey on humans in modern times, such incidents are rare. However, attacks on humans by spotted hyenas are likely to be underreported.Begg, Colleen; Begg, Keith; Muemedi, Oscar (2007
Preliminary data on human carnivore conflict in Niassa National Reserve, Mozambique, particularly fatalities due to lion, spotted hyaena and crocodile
, ''SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique)''
Man-eating spotted hyenas tend to be very large specimens; a pair of man-eating hyenas, responsible for killing 27 people in Mlanje,
Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northe ...
, in 1962, were weighed at after being shot.Kruuk, Hans (2002) ''Hunter and hunted: relationships between carnivores and people'' Cambridge University Press, pp. 64–65, Victims of spotted hyenas tend to be women, children and sick or infirm men, and there are numerous cases of biologists in Africa being forced up trees to escape them. Attacks occur most commonly in September, when many people sleep outdoors, and
bush fire A wildfire, forest fire, bushfire, wildland fire or rural fire is an unplanned, uncontrolled and unpredictable fire in an area of combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identif ...
s make the hunting of wild game difficult for hyenas. In 1903, Hector Duff wrote of how spotted hyenas in the Mzimba district of Angoniland would wait at dawn outside people's huts and attack them when they opened their doors. In 1908–09 in Uganda, spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps.Roosevelt, Theodore (1910) ''African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist'', New York, C. Scribner's sons Spotted hyenas are widely feared in Malawi, where they have been known to occasionally attack people at night, particularly during the hot season when people sleep outside. Hyena attacks were widely reported in Malawi's Phalombe plain, to the north of Michesi Mountain. Five deaths were recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961 when eight people were killed. During the 1960s, Flying Doctors received over two dozen cases of hyena attacks on humans in Kenya. An anecdotal 2004 news report from the
World Wide Fund for Nature The World Wide Fund for Nature Inc. (WWF) is an international non-governmental organization founded in 1961 that works in the field of wilderness preservation and the reduction of human impact on the environment. It was formerly named the W ...
indicates that 35 people were killed by spotted hyenas during a 12-month period in Mozambique along a 20 km stretch of road near the Tanzanian border. Although attacks against living humans are rare, the spotted hyena readily feeds on human corpses. In the tradition of the Maasai and the Hadza, corpses are left in the open for spotted hyenas to eat. A corpse rejected by hyenas is seen as having something wrong with it, and liable to cause social disgrace, therefore it is not uncommon for bodies to be covered in fat and blood from a slaughtered ox. In Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coup and the
Red Terror The Red Terror (russian: Красный террор, krasnyj terror) in Soviet Russia was a campaign of political repression and executions carried out by the Bolsheviks, chiefly through the Cheka, the Bolshevik secret police. It started in ...
. Hyenas habituated to scavenging on human corpses may develop bold behaviours towards living people; hyena attacks on people in southern Sudan increased during the Second Sudanese Civil War, when human corpses were readily available to them.


Urban hyenas

In some parts of Africa, spotted hyenas have begun to frequent metropolitan areas, where groups or "clans" of the animals have become a menace. The Ethiopian capital
Addis Ababa Addis Ababa (; am, አዲስ አበባ, , new flower ; also known as , lit. "natural spring" in Oromo), is the capital and largest city of Ethiopia. It is also served as major administrative center of the Oromia Region. In the 2007 census, t ...
is estimated to have up to a thousand resident hyenas which survive by scavenging rubbish tips and preying on feral dogs and cats. There have also been attacks on homeless people. In 2013, a baby boy was killed by hyenas after being snatched from his mother as she camped near the
Hilton Hotel Hilton Hotels & Resorts (formerly known as Hilton Hotels) is a global brand of full-service hotels and resorts and the flagship brand of American multinational hospitality company Hilton. The original company was founded by Conrad Hilton. As ...
. Some 40 of the animals were reportedly seen alongside a fence bordering the British Embassy compound. In December 2013, a cull was organised and marksmen killed ten hyenas which had occupied wasteland near the city centre.


Hunting and use in traditional medicine

The spotted hyena has been hunted for its body parts for use in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
, for amusement, and for sport, though this is rare, as the species is generally not considered attractive. There is fossil evidence of humans in Middle Pleistocene Europe butchering and presumably consuming spotted hyenas. Such incidences are rare in modern Africa, where most tribes, even those known to eat unusual kinds of meat, generally despise hyena flesh. Several authors during the
Scramble for Africa The Scramble for Africa, also called the Partition of Africa, or Conquest of Africa, was the invasion, annexation, division, and colonization of most of Africa by seven Western European powers during a short period known as New Imperialism ...
attested that, despite its physical strength, the spotted hyena poses no danger to hunters when captured or cornered. It was often the case that native skinners refused to even touch hyena carcasses, though this was not usually a problem, as hyena skins were not considered attractive. In Burkina Faso, the hyena's tail is used for medicinal and magical purposes. In Cameroon, Côte d'Ivoire and Senegal, the animals' whole body is harvested for bushmeat and medicine. In Malawi and Tanzania, the genitalia, nose tips and tails are used for traditional medicine. In Mozambique, traditional healers use various spotted hyena body parts, particularly the paws. Oromo hunters typically go through ritual purification after killing hyenas.Uhlig, Siegbert (2007
''Encyclopaedia Aethiopica: He-N''
Volume 3 of Encyclopaedia Aethiopica, Otto Harrassowitz Verlag, . p. 97.
Kujamaat hunters traditionally treat the spotted hyenas they kill with the same respect due to deceased tribal elders, in order to avoid retribution from hyena spirits acting on behalf of the dead animal. During the early years of Dutch colonisation in southern Africa, hyenas (referred to as "wolves" by the colonists) were especially susceptible to trapping, as their predilection for eating carrion, and lack of caution about enclosed spaces, worked against them. A feature of many frontier farms was the ''wolwehok'' (hyena trap), which was roughly constructed from stone or wood and baited with meat. The trap featured a trap-door, which was designed to shut once the bait was disturbed. In the
Cape Colony The Cape Colony ( nl, Kaapkolonie), also known as the Cape of Good Hope, was a British colony in present-day South Africa named after the Cape of Good Hope, which existed from 1795 to 1802, and again from 1806 to 1910, when it united with ...
, spotted hyenas were often hunted by tracking them to their dens and shooting them as they escaped. Another hunting method was to trap them in their dens and dazzle them with torchlight, before stabbing them in the heart with a long knife.Moodie, John Wedderburn Dunbar (1835) ''Ten Years in South Africa'', Vol. 1, Richard Bentley When chased by
hunting dog A hunting dog is a canine that hunts with or for hunters. There are several different types of hunting dog developed for various tasks and purposes. The major categories of hunting dog include hounds, terriers, dachshunds, cur type dogs, and gu ...
s, spotted hyenas often attack back, unless the dogs are of exceptionally large, powerful breeds.
James Stevenson-Hamilton James Stevenson-Hamilton (2 October 1867 – 10 December 1957) served from 1902–1946 as the first warden of South Africa's Sabi Nature Reserve, which was expanded under his watch and became Kruger National Park in 1926. The Tsonga people ni ...
wrote that wounded spotted hyenas could be dangerous adversaries for hunting dogs, recording an incident in which a hyena managed to kill a dog with a single bite to the neck without breaking the skin. Further difficulties in killing spotted hyenas with dogs include the species' thick skin, which prevents dogs from inflicting serious damage to the animal's muscles.


Spotted hyenas in captivity and as pets

From a husbandry point of view, hyenas are easily kept, as they have few disease problems and it is not uncommon for captive hyenas to reach 15–20 years of age. One study of the hyena immune system showed that captive hyenas had lower levels for immune defenses than hyenas from the wild population that was used to establish the captive population. Nevertheless, the spotted hyena was historically scantily represented in
zoo A zoo (short for zoological garden; also called an animal park or menagerie) is a facility in which animals are kept within enclosures for public exhibition and often bred for conservation purposes. The term ''zoological garden'' refers to z ...
s, and was typically obtained in order to fill empty cages until a more prestigious species could be obtained. In subsequent years, animals considered to be more charismatic were allocated larger and better quality facilities, while hyenas were often relegated to inferior exhibits. In modern times, the species faces spatial competition from more popular animals, especially large canids. Also, many captive individuals have not been closely examined to confirm their sexes, thus resulting in non-breeding pairs often turning out to be same-sexed individuals. As a result, many captive hyena populations are facing extinction. During the 19th century, the species was frequently displayed in travelling
circus A circus is a company of performers who put on diverse entertainment shows that may include clowns, acrobats, trained animals, trapeze acts, musicians, dancers, hoopers, tightrope walkers, jugglers, magicians, ventriloquists, and unicyclis ...
es as oddities. Alfred Brehm wrote that the spotted hyena is harder to tame than the striped hyena, and that performing specimens in circuses were not up to standard.
Sir John Barrow Sir John Barrow, 1st Baronet, (19 June 1764 – 23 November 1848) was an English geographer, linguist, writer and civil servant best known for term as the Second Secretary to the Admiralty from 1804 until 1845. Early life Barrow was born ...
described how spotted hyenas in Sneeuberge were trained to hunt game, writing that they were "as faithful and diligent as any of the common domestic dogs". In Tanzania, spotted hyena cubs may be taken from a communal den by
witchdoctor A witch doctor (also spelled witch-doctor) was originally a type of healer who treated ailments believed to be caused by witchcraft. The term is now more commonly used to refer to healers, particularly in regions which use traditional healing ...
s, in order to increase their social status. An April 2004 BBC article described how a shepherd living in the small town of Qabri Bayah about 50 kilometres from Jigjiga, Ethiopia managed to use a male spotted hyena as a livestock guardian dog, suppressing its urge to leave and find a mate by feeding it special herbs. If not raised with adult members of their kind, captive spotted hyenas will exhibit scent marking behaviours much later in life than wild specimens. Although easily tamed, spotted hyenas are exceedingly difficult to house train, and can be very destructive; a captive, otherwise perfectly tame, specimen in the
Tower of London The Tower of London, officially His Majesty's Royal Palace and Fortress of the Tower of London, is a historic castle on the north bank of the River Thames in central London. It lies within the London Borough of Tower Hamlets, which is sep ...
managed to tear an long plank nailed to its recently repaired enclosure floor with no apparent effort. During the research leading to the composition of his monograph ''The Spotted Hyena: A Study of Predation and Social Behavior,'' Hans Kruuk kept a tame hyena he named Solomon. Kruuk found Solomon's company so congenial, he would have kept him, but Solomon had an insatiable taste for "cheese in the bar of the tourist lounge and bacon off the Chief Park Warden's breakfast table", and no door could hold him back, so Solomon was obliged to live out his days in the Edinburgh Zoo.Kruuk, Hans (1975) ''Hyaena.'' Oxford University Press, London


References


Notes


Bibliography

* * * * * * * * * * * * * * *


Further reading

*Hugo Van Lawick and Jane Goodall. ''Innocent Killers''. Houghton Mifflin Company Boston, 1971 *Mills, M.G.L. ''Kalahari Hyenas: Comparative Behavioral Ecology of Two Species''. The Blackburn Press, 2003


External links


The IUCN Hyaenidae Specialist Group page on spotted hyena
* Davis, Delbert Dwight & Story, Harriette Elizabeth (1949),
The Female External Genitalia of the Spotted Hyena
', Fieldiana: Zoology, vol. 31, no. 33, Chicago Natural History Museum * McPherson, Jami
The Good, the Bad and the Hyena
''BBC Wildlife'', Summer 2008.
Spotted Hyena - Female Genitalia
Check123 Video Encyclopedia {{Good article spotted hyena Carnivorans of Africa Mammals of Sub-Saharan Africa spotted hyena Extant Piacenzian first appearances Apex predators