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    ENGLISH

    SNAMA : KADEK RAY

    KELAS : XI IA 2

    SENIOR HIGH SCHOOL 4 KENDARI

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    HYENA

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    Hyena

    Hyenas (from Greek "" - huaina) are theanimals of the family Hyaenidae,a feliform suborder of the Carnivora. It is the

    smallest biological family in the Carnivora(consisting of four species), and one of thesmallest in the mammalia. Despite their lowdiversity, hyenas are unique and vital

    components to most African andsome Asian ecosystems.

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    Although phylogenetically close

    to felines and viverrids, hyenas are behaviourally

    and morphologically similar to canines in severalaspects (see Convergent evolution) ; both hyenas

    and canines are non-arboreal, cursorial hunters

    which catch prey with their teeth rather than their

    claws. Both eat their food quickly and may store it,

    and their calloused feet with large, blunt, non-

    retractible nails are made for running and making

    sharp turns. However, the hyenas' grooming, scentmarking, defecating habits, mating and parental

    behaviour are consistent with the behaviour of

    other feliforms.

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    Although long reputed to be cowardly

    scavengers, hyenas, especially spotted hyenas,

    do occasionally hunt for themselves, and have

    been known to drive off leopards or lionesses

    from their kills. Hyenas are primarily nocturnalanimals, but may venture from their lairs in the

    early morning hours. With the exception of the

    highly social spotted hyena, hyenas are generallynot gregarious animals, though they may live in

    family groups and congregate at kills.

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    Hyenas first arose in Eurasia during the Miocene period

    from viverrid-like ancestors, and developed into two

    distinct branches ; the lightly built dog-like hyenas andthe robust bone-crushing hyenas. Although the dog-like

    hyenas thrived 15 million years ago (with

    one taxon having colonised North America), they died

    out after a change in climate along with the arrival

    of canids into Eurasia. Of the dog-like hyena lineage,

    only the insectivorous aardwolf survived, while the

    bone-crushing hyenas (whose extant members arethe spotted, brown and striped hyena) became the

    undisputed top scavengers of Eurasia and Africa.

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    Hyenas feature prominently in the folklore and

    mythology of human cultures they are sympatric

    with. Hyenas are mostly viewed with fear and

    contempt, as well as being associated with

    witchcraft, as their body parts are used as

    ingredients in traditional medicine. Among the

    beliefs held by some cultures, hyenas are

    thought to influence peoples spirits, rob graves,and steal livestock and children

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    OriginsHyenas originated in the jungles of Miocene Eurasia 22

    million years ago, when most early feliform species

    were still largely arboreal. The first ancestral hyenas

    were likely similar to the modern banded palm civet ;

    one of the earliest hyena speciesexhumed, Plioviverrops was a lithe, civet-like animal

    which inhabited Eurasia 20-22 millions years ago, and is

    identifiable as a hyaenid by the structure of the middle

    ear and dentition. The lineageof Plioviverrops prospered, and gave rise to

    descendants with longer legs and more pointed jaws, a

    direction similar to that taken by canids in North

    America.

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    Skull ofIctitherium viverrinum, one of the "dog-like"hyenas. American Museum of Natural History

    http://en.wikipedia.org/wiki/File:Ictitherium_viverrinum.JPG
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    Rise and fall of the

    dog-like hyenas

    The descendants

    ofPlioviverrops reached their

    peak 15 million years ago,with more than 30 species

    having been identified.

    Unlike most modern hyena

    species which are specialisedbone-crushers, these dog-like

    hyenas were nimble-bodied,

    wolfish animals; one speciesamong them

    as Ictitherium viverrinum,

    which was similar to a jackal.

    http://en.wikipedia.org/wiki/File:Stripedspottedhyenas.jpg
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    The dog-like hyenas were very numerous; in some

    Miocene fossil sites, the remains ofIctitherium and other

    dog-like hyenas outnumber those of all other carnivores

    combined. The decline of the dog-like hyenas began 5-7

    million years ago during a period of climate change,

    which was exacerbated when canids crossed the Bering

    land bridge to Eurasia. One species, Chasmaporthetes

    ossifragus, managed to cross the land bridge into North

    America, being the only hyena to do

    so. Chasmopothertes managed to survive for some time

    in North America by deviating from the cursorial andbone-crushing niches monopolised by canids, and

    developing into a cheetah-like sprinter. The dog-like

    hyenas died off 1.5 million years ago.

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    Bone-crushing hyenas

    By 10-12 million years ago, the hyena family hadsplit into two distinct groups ; that of the dog-likehyenas and the bone-crushing hyenas. The arrivalof the ancestral bone-crushing hyenas coincidedwith the decline of the similarly built butunrelated Percrocutidae family. The bone-crushing hyenas survived the devastating changesin climate and the arrival of canids which wipedout the dog-like hyenas, though they never

    crossed into North America, as their niche therehad already been taken bythe Borophaginae family.

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    By 5 million years ago, the bone-crushing hyenas

    became the dominant scavengers of Eurasia,primarily feeding on large herbivore carcasses

    felled by sabre-toothed cats . One

    genus, Pachycrocuta, was a 200 kg (440 lb)mega-scavenger which could splinter the bones

    of elephants. With the decline of large

    herbivores by the late ice age, Pachycrocuta was

    replaced by the smaller Crocuta.

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    Rise of modern hyenas

    The aardwolf can trace its lineage directly backto Plioviverrops 15 million years ago, and is theonly survivor of the dog-like hyena lineage. Itssuccess is partly attributed to its insectivorous

    diet, for which it faced no competition fromcanids crossing from North America. Itsunrivaled ability to digest the terpene excretionsfrom soldiertermites is likely a modification of

    the strong digestive system its ancestors used todigest foetid carrion.

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    The striped hyena may have evolved from H.

    namaquensis of Pliocene Africa. Striped hyena fossils are

    common in Africa, with records going back as far as theMiddle Pleistocene and even to the Villafranchian. As

    fossil striped hyenas are absent from

    the Mediterranean region, it is likely that the species is a

    relatively late invader to Eurasia, having likely spreadoutside Africa only after the extinction of spotted

    hyenas in Asia at the end of the Ice Age. The striped

    hyena occurred for some time in Europe during the

    Pleistocene, having been particularly widespreadin France and Germany. It also occurred

    in Montmaurin, Hollabrunn in Austria, the Furninha Cave

    in Portugal and the Genista Caves in Gibraltar.

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    The European form was similar in appearance to

    modern populations, but was larger, being comparable

    in size to the brown hyena.

    The spotted hyena diverged from the striped and

    brown hyena 10 million years ago. Its direct ancestor

    was the Indian Crocuta sivalensis, which lived during

    the Villafranchian. Ancestral spotted hyenas probablydeveloped social behaviours in response to increased

    pressure from rivals on carcasses, thus forcing them to

    operate in teams. Spotted hyenas evolved sharp

    carnassials behind their crushing premolars, thereforethey did not need to wait for their prey to die, as is the

    case for brown and striped hyenas, and thus became

    pack hunters as well as scavengers.

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    They began forming increasingly larger

    territories, necessitated by the fact that their prey

    was often migratory, and long chases in a smallterritory would have caused them to encroach

    into another clan's turf. Spotted hyenas spread

    from their original homeland during the MiddlePleistocene, and quickly colonised a very wide

    area from Europe, to southern Africa

    and China. With the decline of grasslands 12,500

    years ago, Europe experienced a massive loss oflowland habitats favoured by spotted hyenas, and

    a corresponding increase in mixed woodlands.

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    Genera of the Hyaenidae (extinct and

    recent)

    ReconstructionofPachycrocuta breviostris

    A spotted hyena of subfamilyHyaeninae

    http://en.wikipedia.org/wiki/File:Crocuta_crocuta.jpghttp://en.wikipedia.org/wiki/File:Short-faced_hyena1.JPG
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    Their pelage is sparse and coarse with poorly

    developed or absent underfur. Most species have

    a rich mane of long hair running from the withers

    or from the head. With the exception of the

    spotted hyena, hyaenids have striped coats which

    they likely inherited from their vivverid ancestors.

    Their ears are large and have simple basal ridgesand no marginal bursa. Their vertebral column,

    including the cervical region are of limited

    mobility. Hyenas have no baculum. Hyenas havean additional pair of ribs than canids, and their

    tongues are rough like those of felids and

    vivverids.

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    Males in most hyena species are larger than

    females, though the spotted hyena is exceptional,

    as it is the female of the species that outweighsand dominates the male. Also, unlike other

    hyenas, the female spotted hyena's external

    genitalia closely resembles that of the male.

    Their dentition is similar to that of the Felidae, but

    is more specialised for consuming coarse food and

    crushing bones. The carnassials, especially the

    upper, are very powerful and are shifted far backto the point of exertion of peak pressure on the

    jaws.

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    The other teeth, save for the underdeveloped

    upper molars, are powerful, with broad bases and

    cutting edges. The canines are short, but thick androbust. 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. The strength of their jaws is such that both

    striped and spotted hyenas have been recorded tokill dogs with a single bite to the neck without

    breaking the skin.

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    Hyenas lack perinial scent glands, but have a large

    pouch of naked skin located at the anal opening.

    Large anal glands open into it from above the anus.Several sebaceous glands are present between the

    openings of the anal glands and above them. These

    glands produce a white, creamy secretion which is

    pasted onto grass stalks. The odour of this secretion

    is very strong, smelling of boiling cheap soap or

    burning, and can be detected by humans several

    metres downwind. The secretions are primarilyused for territorial marking, though both the

    aardwolf and the striped hyena will spray them

    when attacked.

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    Behaviour

    Hyenas groom themselves often like felids and viverrids,and their way of licking their genitals is very cat-like(sitting on the lower back, legs spread with one legpointing vertically upward). However, unlike otherfeliforms, they do not "wash" their faces. They defecate in

    the same manner as other Carnivora, though they neverraise their legs as canids do when urinating, as urinationserves no territorial function for them. Instead, hyenasmark their territories using their anal glands, a trait found

    also in viverrids and mustelids, but not canids andfelids. When attacked by lions or dogs, striped and brownhyenas will feign death, though the spotted hyena willdefend itself ferociously.

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    The spotted hyena is very vocal, producing a

    number of different sounds consisting of whoops,

    grunts, groans, lows, giggles, yells, growls, laughs and

    whines. The striped hyena is comparatively silent, its

    vocalisations being limited to a chattering laugh and

    howling.

    Mating between hyenas involves a number ofshort copulations with brief intervals, unlike with

    canids in which there is a single, drawn out

    copulation. Spotted hyena cubs are born almost fully

    developed, with their eyes open and erupting incisorsand canines, though lacking adult markings. In

    contrast, striped hyena cubs are born with adult

    markings, closed eyes and small ears.