Fossil range: 26–0 Ma Early Miocene-recent
|Brown hyena (Parahyaena brunnea)|
The Hyaenidae (pronounced /haɪˈɛnɨdiː/) is a mammalian family of order Carnivora. The Hyaenidae family, native to both African and Asian continents, consists of four living species, the Striped Hyena and Brown Hyena (genus Hyaena), the Spotted Hyena (genus Crocuta), and the Aardwolf (genus Proteles).
Hyenas seem to have originated 26 million years ago from arboreal ancestors bearing similarities to the modern Banded Palm Civet. Plioviverrops, one of the earliest hyenas, was a lithe civet-like creature that inhabited Eurasia 20-22 million years ago. Details from the middle ear and dental structure marked it as a primitive hyena. This genus proved successful, its descendants flourishing with more pointed jowls and racier legs, much as the Canidae had done in North America.
Fifteen million years ago, dog-like hyenas flourished, with 30 different species being identified. Unlike some of their modern descendants, these hyenas were not specialized bone-crushers, but were more nimble, wolf-like animals. The dog-like hyenas had canid-like molars, allowing them to supplement their carnivorous diet with vegetation and invertebrates.
Five to seven million years ago, the hyenas were outcompeted by canids traveling from North America to Eurasia via the Bering land bridge.The ancestral aardwolves survived by having adapted themselves to an insectivorous diet to which few canids had specialized. Some hyenas evolved bone-crushing teeth, which allowed them to avoid competition with the canids, resulting in the hyenas eventually outcompeting a family of similarly built bone-crushers called "percrocutoids". The percrocutoids became extinct 7 million years ago, coinciding exactly with the rise of bone-crushing hyena species.
Unlike the canids who flourished in the newly colonized Eurasian continent, only one hyena species, the cheetah-like Chasmaporthetes, managed to cross to North America. It became extinct 1.5 million years ago.
The peak diversity of the Hyenidae was during the Pleistocene, with 4 genera and 9 species of hyena. The bone-crushing hyenas became the Old World's dominant scavengers, managing to take advantage of the amount of meat left over from the kills of sabre-toothed cats. One such species was Pachycrocuta, a 200 kg (440 lb) mega-scavenger that could crush elephant bones. As the sabre-toothed cats began to die out and be replaced by short-fanged felids that were more efficient eaters, more hyenas began to hunt for themselves and began evolving into new species, the modern Spotted Hyena being among them.
The list follows McKenna and Bells Classification of Mammals for prehistoric genera (1997) and Wozencraft (2005) in Wilson and Reeders Mammal Species of the World for extant genera. The Percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family Percrocutidae. Furthermore, the genus Paracrocuta, to which the living brown hyena belongs, is not included into the genus Pachycrocuta, but in the genus Hyaena. The Protelinae (Aardwolves) are not treated as a separate subfamily, but included in the Hyaeninae.
Although hyenas bear some physical resemblance to canids, they make up a separate biological family that is most closely related to Herpestidae (the family of mongooses and meerkats), thereby falling within the Feliformia. All species have a distinctly bear-like gait, due to their front legs being longer than their back legs. The Aardwolf, Striped Hyena, and Brown Hyena have striped pelts and manes lining the top of their necks which erect when frightened. The Spotted Hyena's fur is considerably shorter and spotted rather than striped.
Spotted Hyenas and, to a lesser extent, Striped and Brown Hyenas, have powerful carnassial teeth adapted for cutting flesh and premolars for crushing bone. Spotted Hyenas have a strong bite proportional to their size, but the view that they have the strongest bite is a myth; and a number of other animals (including the Tasmanian devil) are proportionately stronger. The Aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dentition as the other three species. The dental formula for all hyena species is:
Labiolingually, their mandibles are much stronger at the canine teeth than in canids, reflecting the fact that hyenas crack bones with both their anterior dentition and premolars, unlike canids which do so with their post-carnassial molars. Like felids, hyenas lack the rearward molars of canids and vivverids. By organising their teeth so that the bone-crushing premolars do not interfere with the meat-slicing carnassials to the rear, hyenas can crush bone without blunting the carnassials' blades.
Spotted Hyena societies are more complex than those of other carnivorous mammals and have been reported to be remarkably similar to those of cercopithecine primates in respect to group size, structure, competition, and cooperation. One indication of hyena intelligence is that they will move their killed prey closer together to protect them from scavengers. Another indication is their strategic hunting methods.
The majority of hyena species show little sexual dimorphism, with males being only slightly larger than the females. The Spotted Hyena is an exception to this, with females larger than males. One unusual feature of the Spotted Hyena is that females have an enlarged clitoris, called a pseudo-penis, demi-penis, or sometimes mistakenly referred to as a nanophallus. Female hyenas give birth, copulate, and urinate through their protruding genitalia, which stretches to allow the male penis to enter for copulation; it also stretches during birth. The anatomical position of the genitalia gives females complete control over which males are allowed to mate with them.
Researchers originally thought that one cause of this characteristic of the genitals was androgens that were introduced to the fetus very early on in its development. However, it was discovered that when the androgens were held back from the female fetus, the development of her genitalia was not altered. Spotted Hyenas have a matriarchal social structure that some biologists speculate evolved because it is in the best interests of the female hyena to dominate the male hyena as it provides no assistance in rearing the cubs.
All species excrete an oily, yellow substance from their anal glands onto objects to mark their territories. When scent marking the anal pouch is turned inside out, or everted. Hyenas also do this as a submissive posture to more dominant hyenas. Genitals, the anal area and the anal glands are sniffed during greeting behavior in which each hyena lifts its leg and allows the other to sniff its anal sacks and genitals. All four species maintain latrines far from the main denning area where dung is deposited. Scent marking is also done by scraping the ground with the paws, which deposits scent from glands on the bottoms of the feet. Hyenas do not raise their legs when urinating as male or dominant canids do.
Three of the four species of hyena are restricted to sub-Saharan Africa, where they live in drier environments such as savannah, bushland and desert. The fourth species, the Striped Hyena, is found in northern and eastern Africa as well as in Asia from the Middle East to India.
Except for the aardwolf, all living hyena species are hunters and scavengers. They have extremely strong jaws in relation to their body size and have a very powerful digestive system with highly acidic fluids, making them capable of eating and digesting their entire prey, including skin, teeth, horns and bones. Hair and hooves are usually regurgitated. Because their digestive system deals very well with bacteria, they have no aversion to and readily eat carrion.
The Spotted Hyena is primarily a predator, unlike some of its cousins. Spotted Hyenas are successful pack hunters of small to large sized ungulates, and are the most abundant carnivores in Sub-Saharan Africa.
Many cultures, including those in Africa, have historically viewed the hyena negatively, associating them with gluttony, uncleanliness and cowardice.
Part of their bad reputation may stem from the hyena's tendency to scavenge graves for food. They are one of the few creatures naturally suited for this, due to their ability to devour and digest every part of a carcass, including bone. The word hyena is derived from the Greek hyaina, meaning "pig", and has a long association with cruelty, treachery and greed.
In Malawi, in the local language large hyenas reputed to be a man-eater were called lipwereri and the ordinary hyena was called a fisi. The Bouda is a mythical tribe reputed to house members able to transform into hyenas. Belief in "Werehyenas" is so entrenched within the traditional lore of the Bornu people of north-eastern Nigeria, that their language even contains a special word, bultungin, which translates as "I change myself into a hyena".
The haunting laughter-like calls of the Spotted Hyena inspired the idea in local cultures that they could imitate human voices and call their victims by name. Hyenas are also associated with divination and sometimes thought of as tools of demons and witches. In African folklore, witches and sorcerers are thought to ride hyenas or even turn into them.
Early naturalists thought hyenas were hermaphrodites or commonly practiced homosexuality, largely due to the female Spotted Hyena's unique urogenital system. According to early writings such as Ovid's Metamorphoses and the Physiologus, the hyena continually changed its sex and nature from male to female and back again. In Paedagogus, Clement of Alexandria noted that the hyena (along with the hare) was "quite obsessed with sexual intercourse." Many Europeans associated the hyena with sexual deformity, prostitution and deviant sexual behavior.
Hyenas (usually "Laughing Hyenas") have been used in animated movies many times as well as having been rendered in live action films, commonly cast as hysterical and unhinged villains. Examples include Shenzi, Banzai and Ed from the Disney animated film The Lion King, one ball-playing individual in Bedknobs and Broomsticks, Harley Quinn's pets, Bud and Lou, and the laughing Hyena in The Lady and the Tramp.
HYENA, a name applicable to all the representatives of the mammalian family Hyaenidae, a group of Carnivora allied to the civets. From all other large Carnivora except the African hunting-dog, hyenas are distinguished by having only four toes on each foot, and are further characterized by the length of the fore-legs as compared with the hind pair, the non-retractile claws, and the enormous strength of the jaws and teeth, which enables them to break the hardest bones and to retain what they have seized with unrelaxing grip.
I See further under Scyphomedusae.
r The striped hyena (Hyaena striata) is the most widely distributed species, being found throughout India, Persia, Asia Minor, and North and East Africa, the East African form constituting a distinct, race, H. striata schillingsi; while there are also several distinct Asiatic races. The species resembles a wolf in size, and is greyish-brown in colour, marked with indistinct longitudinal stripes of a darker hue, while the legs are transversely striped. The hairs on the body are long, especially on the ridge of the neck and back, where they form a distinct mane, which is continued along the tail. Nocturnal in habits, FIG. 1. - The Striped Hyena (Hyaena striata). it prefers by day the gloom of caves and ruins, or of the burrows which it occasionally forms, and issues forth at sunset, when it commences its unearthly howling. When the animal is excited, the howl changes into what has been compared to demoniac laughter, whence the name of "laughing-hyena." These creatures feed chiefly on carrion, and thus perform useful service by devouring remains which might otherwise pollute the air. Even human dead are not safe from their attacks, their powerful claws enabling them to gain access to newly interred bodies in cemeteries. Occasionally (writes Dr W. T. Blanford) sheep or FIG. 2. - The Spotted Hyena (Hyaena crocuta). goats, and more often dogs, are carried off, and the latter, at all events, are often taken alive to the animal's den. This species appears to be solitary in habits, and it is rare to meet with more than two together. The cowardice of this hyena is proverbial; despite its powerful teeth, it rarely attempts to defend itself. A very different animal is the spotted hyena, Hyaena (Crocuta) crocuta, which has the sectorial teeth of a more cat-like type, and is marked by dark-brown spots on a yellowish ground, while the mane is much less distinct. At the Cape it was formerly common, and occasionally committed great havoc among the cattle, while it did not hesitate to enter the Kaffir dwellings at night and carry off children sleeping by their mothers. By persistent trapping and shooting, its numbers have now been considerably reduced, with the result, however, of making it exceedingly wary, so that it is not readily caught in any trap with which it has had an opportunity of becoming acquainted. Its range extends from Abyssinia to the Cape. The Abyssinian form has been regarded as a distinct species, under the name of H. liontiewi, but this, like various more southern forms, is but regarded as a local race. The brown hyena (H. brunnea) is South African, ranging to Angola on the west and Kilimanjaro on the east. In size it resembles the striped hyena, but differs in appearance, owing to the fringe of long hair covering the neck and fore part of the back. The general hue is ashy-brown, with the hair lighter on the neck (forming a collar), chest and belly; while the legs are banded with dark brown. This species is not often seen, as it remains concealed during the day. Those frequenting the coast feed on dead fish, crabs and an occasional stranded whale, though they are also a danger to the sheep and cattle kraal. Strand-wolf is the local name at the Cape.
Although hyenas are now confined to the warmer regions of the Old World, fossil remains show that they had a more northerly range during Tertiary times; the European cave-hyena being a form of the spotted species, known as H. crocuta spelaea. Fossil hyenas occur in the Lower Pliocene of Greece, China, India, &c.; while remains indistinguishable from those of the striped species have been found in the Upper Pliocene of England and Italy.
Hyeres, a town in the department of the Var in S.E. France, I I m. by rail E. of Toulon. In 1906 the population of the commune was 17,790, of the town 10,464; the population of the former was more than doubled in the last decade of the 19th century. Hyeres is celebrated (as is also its fashionable suburb, Costebelle, nearer the seashore) as a winter health resort. The town proper is situated about 22 m. from the seashore, and on the southwestern slope of a steep hill (669 ft., belonging to the Maurettes chain, 961 ft.), which is one of the westernmost spurs of the thickly wooded Montagnes des Maures. It is sheltered from the north-east and east winds, but is exposed to the cold north-west wind or mistral. Towards the south and south-east a fertile plain, once famous for its orange groves, but now mainly covered by vineyards and farms, stretches to the sea, while to the southwest, across a narrow valley, rises a cluster of low hills, on which is the suburb of Costebelle. The older portion of the town is still surrounded, on the north and east, by its ancient, though dilapidated medieval walls, and is a labyrinth of steep and dirty streets. The more modern quarter which has grown up at the southern foot of the hill has handsome broad boulevards and villas, many of them with beautiful gardens, filled with semis tropical plants. Among the objects of interest in the old town are: the house (Rue Rabaton, 7) where B. Massillon (1663-1742), the famous pulpit orator, was born; the parish church of St Louis, built originally in the 13th century by the Cordelier or Franciscan friars, but completely restored in the earlier part of the 19th century; and the site of the old château, on the summit of the hill, now occupied by a villa. The plain between the new town and the sea is occupied by large nurseries, an excellent jardin d'acclimatation, and many market gardens, which supply Paris and London with early fruits and vegetables, especially artichokes, as well as with roses in winter. There are extensive salt beds (salines) both on the peninsula of Giens, S. of the town, and also E. of the town. To the east of the Giens peninsula is the fine natural harbour of Hyeres, as well as three thinly populated islands (the Stoechades of the ancients), Porquerolles, Port Cros and Le Levant, which are grouped together under the common name of Iles d'Hyeres.
The town of Hyeres seems to have been founded in the 10th century, as a place of defence against pirates, and takes its name from the aires (hierbo in the Provencal dialect), or threshingfloors for corn, which then occupied its site. It passed from the possession of the viscounts of Marseilles to Charles of Anjou, count of Provence, and brother of St Louis (the latter landed here in 1254, on his return from Egypt). The château was.
dismantled by Henri IV., but thanks to its walls, the town resisted in 1707 an attack made by the duke of Savoy.
See Ch. Lentheric, La Provence Maritime ancienne et moderne (chap. 5) (Paris, 1880). (W. A. B. C.)