Hyenas

Hyena is a redirect to this article. Further meanings are listed under hyena (disambiguation) .

Hyenas
Spotted hyena ( Crocuta crocuta )
Systematics
Class : Mammals (mammalia) Subclass : Higher mammals (Eutheria) Superordinate : Laurasiatheria Order : Predators (Carnivora) Subordination : Feline (Feliformia) Family : Hyenas
Scientific name
Hyaenidae
Gray , 1821

The aardwolf is the smallest hyena species and its teeth differ significantly from other species

The hyenas (Hyaenidae) are a mammal family from the order of the carnivores (Carnivora) with four recent species that live in large parts of Africa as well as in western and southern Asia . The term hyena is often used to refer to the largest and most individual species of spotted hyena , whose rivalry with lions is a popular theme in animal films .

There are two subfamilies of hyenas. The three types of the first, the real hyenas (Hyaeninae), are characterized by strong teeth: the spotted hyena , the striped hyena and the saddleback hyena . The spotted hyena feeds primarily on active hunting, while the striped hyena and the black hyena are primarily scavengers . The monotypical second subfamily (Protelinae) is represented by the aardwolf , which feeds almost exclusively on the representatives of a termite genus and whose molars are therefore greatly reduced in size. The main threat to the hyenas is hunting by humans.

features

General build and coat

The head-trunk length is 55-160  cm , the tail is relatively short with 20-40 cm. The shoulder height measures 45–81 cm, the front legs are longer and stronger than the rear legs, which is the reason for the sloping back that is typical of all hyena species. The females of the spotted hyenas, the largest species, are around 10% larger than the males; in the other species there is no significant sex dimorphism in terms of size. The actual hyenas weigh 26–55  kg , with individual spotted hyenas reaching up to 86 kg; At 8-14 kg, the aardwolf is by far the smallest and lightest species. Hyenas have four toes on their front and rear legs, except for the aardwolf, which has one more toe on each front leg. The paws have blunt, non-retractable claws .

The outer hairs are coarse, with the exception of the spotted hyena, all species have a long mane on the back , which extends from the ears to the tail. This mane can be straightened up, making the animal appear larger. The fur of the individual species shows different shades of brown, the spotted hyena is spotted, the striped hyena and the aardwolf are striped, only the white hyena is largely monochrome. The tail is bushy.

The females have one to three pairs of teats , the males - in contrast to most other predators - lack the penis bones (baculum). Female spotted hyenas have a masculinization ("masculinization") that is unique among mammals: the clitoris is enlarged, the labia are closed and form a scrotum-like structure. The position of this "dummy penis" prevents mating without the partner's consent. Urination, copulation, and childbirth take place through the clitoris. Growing striped hyenas have bulges in the genital tract, but when fully grown they, like the other two hyena species, show no special features in the construction of the genital tract. Both sexes have a well-developed anal pouch from which a secretion is released that is used to mark the territory .

Head and teeth

Skull of a spotted hyena: The domed skull and the strong molars are typical of the real hyenas (Hyaeninae)

Diagnostic features of hyenas are found in the sphenoid bone , where the alisphenoid canal is absent, and in the bones of the middle ear region , where the endotympanic part of the tympanic membrane is small but the ectotympanic part is distended. In addition, the structure of the skull and the teeth show the greatest differences between the two subfamilies: The real hyenas have a massive head on their powerful neck and their muzzle is broad, while the aardwolf's head is slender with a pointed snout.

The hyena's incisors are unspecialized, and the outer teeth are larger than the others. The canines are enlarged. The real hyenas have strong teeth. The premolars are adapted and enlarged for bone breaking, especially the third upper and third lower premolars. Your tooth enamel has a complex structure that prevents teeth from breaking. As in all land carnivores , the fourth upper premolar and the first lower molar are developed into fangs ; these teeth are blade-shaped and are used to cut meat. The molars behind the fangs are either reduced in size or completely absent, leaving more space for the remaining molars: the premolars become wider and the fangs are better protected from abrasion. The tooth formula of the real hyenas consists of 3/3 I , 1/1 C , 4/3 P and 1/1 M , a total of 34 teeth. The aardwolf's canine teeth are used exclusively for confrontation with conspecifics. The molars have receded into small, widely spaced pins, the number of which can vary. The aardwolf tooth formula is 3/3 I , 1/1 C , 3 / 1–2 P and 1 / 1–2 M , a total of 28–32 teeth.

Connected to the strong teeth of the real hyenas is a strong chewing muscles; the temporalis muscle has a high crest at the point of attachment to the skull. The domed skull ensures better implementation of the biting forces . Thanks to their extraordinary jaw apparatus, spotted hyenas can develop biting forces of over 9 kN . They are able to break the leg bones of giraffes , rhinos, and hippos that are over 7 cm in diameter . Adaptations of the aardwolves to insect food consist of a broad palate with a broad, spatulate tongue covered with large, cone-shaped papillae .

The eyes of all hyenas are equipped with a tapetum lucidum , which enables good visual performance at night. The ears are large and pointed, only in the spotted hyena they are rounded.

distribution and habitat

Hyenas are native to large parts of Africa and western and southern Asia . In Africa their distribution area extends from the Atlas Mountains to South Africa , but they are absent in the pure desert regions of the Sahara and the Congo Basin . In Asia they occur from Turkey and the Arabian Peninsula via Afghanistan to India . Even in the Pleistocene , they were spread over large parts of Eurasia , with Chasmaporthetes , however, only one extinct genus is known that also occurred in North America. There were never any hyenas in South America or Australia.

In Asia there is only one species with the striped hyena; this also inhabits northern Africa and thus has the northernmost range of all species. The spotted hyena is native to large parts of sub-Saharan Africa. The aardwolf lives in two separate areas in eastern and southern Africa, while the black hyena lives in a relatively small area in the south of the continent.

In general, hyenas inhabit dry areas such as semi-deserts , savannas , bush steppes and rocky mountainous areas; sometimes they can also be found in swamp areas and mountain forests. In the Ethiopian highlands they can be found up to 4100  m . However, they avoid pure sandy deserts as well as lowland rainforests. Hyenas are usually not very picky about their habitat, each of the four species is found in several habitats . They are not shy of people and occasionally stay in the vicinity of human settlements.

Way of life

Movement and activity times

Saddleback hyenas , like all hyenas, are predominantly nocturnal

Hyenas are digitigrad (toe walkers) and only stay on the ground. They cannot climb trees, but they can cope well with rocky terrain. They are very persistent animals: Svalbard hyenas can cover more than 50 kilometers per night. All hyena species are largely nocturnal; they rarely go for food at dusk or - in the case of the spotted hyena - during the day in cloudy, rainy weather. During the day they sleep in earthworks, hidden in the bushes or on the ground. For example, aardwolves often sleep in burrows that they have taken over from spring hares or other animals; sometimes hyenas also build their own hiding places or make do with crevices in the rock. The spotted hyena does not use any burrows other than to raise the young.

Social and territorial behavior

When social behavior , there is a wide variety. Spotted hyenas and spotted hyenas live in groups called "clans". In both species, groups of related females form the core of a clan, the reproductive males have immigrated and are not related to the females. The social behavior of the spotted hyenas is unique among predators : It resembles that of some Old World monkeys , such as baboons . In this species, the clans can include up to 80 animals, which are repeatedly divided into smaller subgroups and come together again. The females are dominant and establish a strict hierarchy, with the ranks being hereditary as the mothers help their daughters achieve the same position as them. The males are always subordinate to the females and their group rank is higher, the longer they belong to the group. The saddleback hyenas clans include 4–14 animals, there is a variance in the way of life depending on the habitat. Only at high population density to establish hierarchies ; Males and females each have their own hierarchies, and both sexes are equal. Various observations have been made in the striped hyena, the least researched species: there have been reports of solitary animals, stable pair bonds and living together in groups. Presumably, the social behavior of this species is variable. The aardwolves enter into stable and long-lasting relationships when rearing their young, but the young were often not conceived by the rearing male. Outside the mating season, aardwolves show hardly any social behavior: They live in separate burrows and go foraging separately.

Striped hyenas , like all hyenas, are territorial animals

Hyenas are territorial animals, the size of the territory depends on the species and the food supply: The territories of the spotted hyenas in the prey rich savannahs of East Africa measure around 20 km², while the territories of spotted hyenas in the arid regions of southern Africa can be over 1000 km² . The ground wolves' territory comprises around 3,000 termite mounds and measures 1.5–4 km². The areas are marked with the strongly smelling secretion of their anal bags , which is whitish or yellowish in color. It is touched on tufts of grass or other objects in a half crouching position. While spotted hyenas often only mark the territorial boundaries, the other species often also add their scent traces inside the territory. In addition, all species create pits near the territorial boundaries or on routes that are frequently traveled, in which they regularly defecate . If a hyena encounters an alien animal in its own territory, it tries to chase it away. With the exception of the spotted hyena, it straightens its back mane and ruffles its tail hair, making it appear larger. If that doesn't help, she tries to drive away the intruder, these hunts end at the border of the territory. But sometimes there are also fights that are fought with bites.

communication

Since the territories are often huge and the animals often travel alone, olfactory communication , i.e. by means of smells, plays an important role. With the help of the anal bag secretion, hyenas can identify the sex, reproductive status and group membership of other hyenas. The real hyenas have a special greeting ritual that members of the same group show when they come together: They sniff the other animal's nose or anal pouch or lick its back. In spotted hyenas, the erect sexual organs - both in males and females - play an additional role, and are sniffed or licked by the animal opposite.

Three of the four hyena species make only a few sounds . At best, they emit growling or screeching noises that can only be heard over short distances. In contrast, the spotted hyena has a rich repertoire of verbal communication. The most commonly heard sound is a loud wuup , which can be heard over several kilometers and is used to contact other clan members. They also make grunting, crying and mooing noises. Finally, the laughing or giggling sound, which is similar to human laughter, is known; it signals that the animal accepts a lower rank.

food

Spotted hyenas eat an impala

The four hyena species have occupied three different ecological niches in terms of nutrition . Spotted hyenas are active hunters who hunt 60 to 95% of their prey themselves. They have a very wide range of prey: the spectrum ranges from insects to elephants. Most often, however, they kill larger ungulates, such as various antelopes - such as wildebeest and gazelles - or zebras . They hunt individually or in groups, depending on the prey. In doing so, they do not sneak up on their victims, but rely on their perseverance . The form of food acquisition is also flexible: In addition to animals that they hunt themselves, they also eat carrion or practice kleptoparasitism , i.e. they hunt other carnivores off their prey. This has been observed in jackals , cheetahs , leopards , African wild dogs , the other two real hyenas and lions .

Striped hyenas and saddleback hyenas are scavengers who also eat prey and plant material that they have killed themselves. The carrion of larger vertebrates makes up a large part of their food . Thanks to their strong teeth, they can also break thick bones or turtle shells, and their efficient digestive system utilizes all parts of an animal's body with the exception of hair, hooves and horns. The bacterial toxins contained in the carrion do not affect your digestive or immune systems. Small mammals, birds and their eggs as well as insects complete their menu. It is unclear to what extent they eat larger prey that they have killed themselves. They are not good hunters, most hunts fail. An exception are the saddleback hyenas of the Namibian coast, which hunt the young South African fur seals living there with great success .

Erdwolves eat almost exclusively termites , and they have specialized in the animals of the Trinervitermes genus . The termites of this genus move in large groups on the surface of the earth at night and are licked by aardwolves with their sticky tongues. The soldiers of these termites secrete a poison that is incompatible with numerous other insectivorous mammals, one exception being the aardvark . Thanks to their insensitivity to this poison, aardwolves largely avoid food competition with other animals.

Apart from the spotted hyenas, which hunt in groups, hyenas usually go for food individually, but in the case of larger prey or carcasses, several hyenas can come together and eat together.

Hyenas don't need to drink; however, when water is available, they drink daily. Striped hyenas and saddleback hyenas cover their fluid needs with cucurbits and other plants.

Reproduction

Young female hyena

The mating behavior of the hyenas is different. Often a promiscuous behavior prevails , that is, males and females reproduce with several partners each. In addition, polyandry is sometimes found in striped hyenas , which is rare in mammals, i.e. a female has several male mating partners. The choice of partner is also variable: In the case of saddleback hyenas, the nomadic males, who roam the area of ​​the clan without their own territory and solitary, are chosen as partners, which may depend on the food supply. In the case of aardwolves, in around 40% of all breedings, it is not the partner with whom the female cohabits that mates, but a more aggressive, stronger male from a neighboring territory. Serengeti Hyena males invest a lot of time in courtship. Gentle males have far more success in mating than aggressive mates.

In most cases, reproduction is not seasonal, but can occur all year round. After a gestation period of around 90 to 110 days , the female gives birth to one to four young, in human care there can be five. The level of development of newborns varies: while the incisors and canines of the deciduous dentition are already present in spotted hyenas at birth and the eyes are open, striped hyenas and saddleback hyenas are less developed and their eyes are still closed. Newborn spotted hyenas are colored black, in the other species the fur color of the young is similar to that of the adult, only instead of the mane on the back there is a dark eel line .

Newborn hyenas spend their first few weeks in a burrow. With spotted hyenas and sometimes also with spotted hyenas, there are community buildings in which the young of a clan grow up together. In the aardwolves' den, the males guard the offspring in their mother's den that were not or only partially conceived by them. As far as is known, this is unique among mammals. After a few weeks, the young animals begin to explore the area outside the den. After a few months, they undertake their first forays in search of food, initially accompanied by a full-grown animal, then later alone. Young hyenas are suckled for a relatively long time, real hyenas are not finally weaned until they are 12-16 months old. Sexual maturity occurs in the second or third year of life .

Hyenas and humans

etymology

The word “hyena” is already attested in Old High German as ijēna , in Middle High German it stands for clarification and later hienna . Ultimately, it comes from the Latin hyaena from the Greek ὕαινα ( hýaina ), which is derived from the word ὗς ( hӯs ), "pig". Presumably, the hyena has been compared to a pig because of its bristle back.

Hyenas in culture

Painting of the fight between a striped hyena and two domestic dogs

There are numerous myths about hyenas in Africa and Asia , and often no distinction is made between the individual species. They play an ambivalent role in African narratives : on the one hand, they are considered cruel and dangerous animals, but sometimes they also symbolize strength and endurance and are considered sacred animals. In the mythology of the tabwa from eastern Africa, a spotted hyena brought the sun to warm the earth, but in West African cultures they symbolize bad qualities. The witches are said to ride these animals, and hyena masks are used in rites of some peoples to convey the cunning and other characteristics to the wearer. The alter ego to the nocturnal, wild hyenas embodies the diurnal, domesticated dogs in their relationship with humans. In the myths of the Beng, an ethnic group in Ivory Coast , bones symbolize the opposing relationship of both animals to humans. Dogs chew bones of wild animals thrown for them by humans (voluntarily), while hyenas are decried by the Beng as grave robbers who dig up human bones (against human will). Because of their magical abilities corresponding to hyenas, dogs are sacrificed to the deity Nya in the south of Mali so that she can take up the fight against the dreaded witchcraft. For the Yoruba , hyenas are considered to be symbols of the end of all things, as they clear away the carcasses left by other carnivores.

In the Near and Middle East, hyenas are considered to be manifestations of jinn . Among the Bedouins there are numerous myths about encounters with mystical hyenas or hyena-human hybrid beings, which are called kaftar . The genitals were thought to be a love spell, the tongue was supposed to be a cure for tumors , and other parts of the body were also believed to have healing powers; this is already known from ancient Greece and ancient Rome . In Afghanistan, fights were held between domestic dogs and hyenas in order to then use the body parts in magical rituals.

Already Aristotle wrote in his work Historia animalium that the hyena loves rotten meat and digs in cemeteries to get something to eat. However, he rejects the claim that hyenas are hermaphrodites , although this error persisted into the 20th century. Also Pliny the Elder deals with the hyenas in his work Naturalis Historia , in which he mainly reflects the insights of Aristotle and describes the healing properties of various body parts. The Physiologus , an early Christian animal symbolism, writes that the hyenas would be able to change their sex. They are therefore compared with deceitful people; this interpretation is also applied in an anti-Semitic way to the Jews, who first worshiped the true God, but later allegedly worshiped idols. This happens in the letter of Barnabas , an early Christian theological treatise directed against the Jews. There, in verse 10.7, the ancient view of the hyenas being two sexes is adopted in order to justify a ban on consumption.

Various bestiaries of the Middle Ages repeat these views. In various more recent works, too, hyenas are often portrayed negatively and the mistake that they are hermaphrodites or that they regularly change their sex is repeated. The bad reputation is still evident, for example, in Brehm's animal life , as Alfred Brehm writes about the spotted hyena: “Among all predatory animals, it is undoubtedly the most misshapen, nasty phenomenon; In addition to this, however, there are also the spiritual qualities to make the animal hateful. ”They are still considered insidious and cowardly scavengers today. For example, an intrusive telephone interview of bank robbers by journalists from the daily newspaper Austria was called "hyena journalism".

See also : hyena people

threat

It happens again and again that hyenas invade pastures and kill domestic animals. Spotted hyenas also occasionally kill people, for example when they sleep outdoors without protection. There are reports of the striped hyena invading cemeteries, digging up corpses, and eating. For these reasons, they are often pursued and hunted down with poison bait, firearms or traps. Another reason for hunting are the healing powers attributed to various parts of the hyena's body. The Tuareg fattened and ate striped hyenas until at least the 1940s, a practice also known from ancient Egypt . Further threats are the destruction of their habitat and the decline in prey animals due to human influence. Automobile traffic represents a hazard. This risk is increased by the fact that hyenas often eat the carcasses of animals run over directly on the street and are careless towards vehicles following behind.

With the exception of the aardwolf, the populations of all hyena species are declining. In the case of aardwolves, the large-scale grazing industry tends to increase their preferred genus of termites, which has a positive effect on the total population. Estimates of the total population of the various hyena species amount to 27,000–47,000 spotted hyenas, 5,000–8,000 saddleback hyenas, 5,000–14,000 striped hyenas and at least several thousand aard wolves. The IUCN lists the striped hyena and the black-tailed hyena as " near threatened " and the aardwolf and the spotted hyena as "not at risk" ( least concern ).

Systematics and tribal history

External system

The mongooses - here a zebra mongoose - and Malagasy predators are the closest living relatives of the hyenas

In spite of their dog-like appearance, the hyenas are classified as cats within the predators , which is secured by skull features, in particular the construction of the tympanic cavity . Relations with the other feline species were long unexplained; hyenas were thought to be close relatives of cats , the mongooses, or an independent, early side branch of the feline species. By molecular studies it was found that the Schwestertaxon the hyenas a common clade from Mangusten and Malagasy predators is. The position of the hyenas within the feline is shown in the following cladogram :

Feline (Feliformia or Feloidea)	Mongooses (Herpestidae)      Malagasy carnivores (Eupleridae)      Hyenas (Hyaenidae)      Crawling cats (Viverridae)      Cats (Felidae)      Linsangs (Prionodontidae)      African palm civet (Nandiniidae)

The lines of development between the hyenas on the one hand and the mongooses and Malagasy carnivores on the other have separated around 29.2 million years ago.

Internal systematics of today's hyenas

The spotted hyena is the sister taxon of a common clade of striped hyena and black horned hyena

There are four recent hyena species:

• Striped hyena ( Hyaena hyaena ( Linnaeus , 1758))
• Saddleback hyena ( Parahyaena brunnea ( Thunberg , 1820); partly also Hyaena brunnea )
• Spotted hyena ( Crocuta crocuta ( Erxleben , 1777))
• Aardwolf ( Proteles cristatus ( Sparrman , 1783))

These four species are divided into two subfamilies, on the one hand the Protelinae with the aardwolf, on the other hand the Hyaeninae (real hyenas) with the other three species. Some systematics consider the differences in physique and way of life between the aardwolf and the real hyenas to be so great that they place the aardwolf in a separate family, the Protelidae. This division is not carried out by more recent taxonomic publications. Molecular studies have confirmed the monophyly , that is, the descent from a common ancestral form, of both the hyenas and the real hyenas. Accordingly, the aardwolf is the sister taxon of the real hyenas and the spotted hyena is the sister taxon of a common clade of striped hyena and saddleback hyena. This becomes clear in the following cladogram:

Hyenas (Hyaenidae)	Actual hyenas (Hyaeninae)    Spotted hyena ( Crocuta crocuta )   NN    Striped hyena ( Hyaena hyaena )      Saddleback hyena ( Parahyaena brunnea )   Protelinae    Aardwolf ( Proteles cristata )

The lines of development between the aardwolf and the real hyenas separated around 10.6 million years ago, that between the spotted hyena and the striped hyena and saddleback hyena around 8.6 million years ago. The split between striped hyenas and saddleback hyenas happened around 4.2 million years ago.

Overview of the recent and fossil genera of the hyenas

The list below of the fossil and recent hyena genera largely follows the classification of Werdelin and Solounias (1991) and McKenna and Bell (1997) for the extinct representatives. The current genera were supplemented by Wozencraft (2005) in Wilson and Reeder's Mammal Species of the World . In contrast to the classification by McKenna and Bell, however , the percrucotids are not listed, but are viewed as a separate family. The extinct African forms received a detailed revision by Werdelin and Peigné in 2010, and further studies were also taken into account:

• Family Hyaenidae Gray , 1821

• † Tongxinictis Werdelin & Solounias , 1991 (Middle Miocene of Asia)
• † Belbus Werdelin & Solounias , 1991 (Upper Miocene of Greece)
• † Allohyaena Kretzoi , 1938 (Upper Miocene of Europe)

• † Subfamily Ictitheriinae Trouessart , 1897

• † Plioviverrops Kretzoi , 1938 (including Jordanictis , Protoviverrops , Mesoviverrops ; Lower Miocene to Lower Pliocene of Europe and Upper Miocene of Asia)
• † Ictitherium Roth & Wagner , 1854 (= Galeotherium ; including Lepthyaena , Sinictitherium , Paraictitherium ; Middle Miocene of Africa, Upper Miocene to early Pliocene of Eurasia)
• † Thalassictis Nordmann , 1950 (including Palhyaena , Miohyaena , Hyaenalopex ; Middle to Upper Miocene of Asia, Upper Miocene of Europe)
• † Hyaenictitherium Zdansky , 1924 (Upper Miocene and Lower Pliocene of North and East Africa)
• † Hyaenotherium Semenov , 1989 (Upper Miocene (to Lower Pliocene?) Eurasia)
• † Miohyaenotherium Semenov , 1989 (Upper Miocene of Europe)
• † Lycyaena Hensel , 1863 (Upper Miocene of Eurasia)
• † Lycyaenops Kretzoi , 1938 (Middle Miocene of North Africa)
• † Tungurictis Colbert , 1939 (Middle Miocene of Asia, possibly also Africa and Europe)
• † Protictitherium Kretzoi , 1938 (Middle Miocene of Africa, Middle Miocene of Asia, Middle to Upper Miocene of Europe)

• Subfamily Hyaeninae Gray , 1821

• † Palinhyaena Qiu, Huang & Guo , 1979 (Upper Miocene of Asia)
• † Ikelohyaena Werdelin & Solounias , 1991 (Lower Pliocene of Africa)
• Hyaena Brisson , 1762 (= Euhyaena , Hyena , Pliohyaena , Anomalopithecus ; early Pliocene (possibly Middle Miocene) to today in Africa, Upper Pliocene (possibly Lower Miocene) to Late Pleistocene in Europe, Upper Pliocene to today in Asia)
• Parahyaena Hendey , 1974 (from the Lower Pliocene fossil in eastern and southern Africa until today)
• † Pliocrocuta Kretzoi , 1938 (Pliocene of Eurasia and North Africa)
• † Hyaenictis Gaudry , 1861 (Upper Miocene of Asia?, Upper Miocene of Europe, Lower Pliocene (possibly also Lower Pleistocene) of Africa)
• † Leecyaena Young & Liu , 1948 (Upper, possibly also Lower Miocene of Asia)
• † Chasmaporthetes Hay , 1921 (= Ailuriaena ; including Euryboas ; Upper Miocene to Lower Pleistocene of Eurasia, Lower Pliocene to Upper Pliocene or Lower Pleistocene of Africa, Upper Pliocene to Early Pleistocene of North America)
• † Pachycrocuta Kretzoi , 1938 (Pliocene and Pleistocene of Eurasia and Africa)
• † Adcrocuta Kretzoi , 1938 (Upper Miocene of Eurasia)
• Crocuta Kaup , 1828 (= Crocotta ; including Eucrocuta ; Upper Pliocene to this day in Africa, Upper Pliocene to Upper Pleistocene in Eurasia)

• Subfamily Protelinae Flower , 1869

• Proteles I. Geoffroy St.-Hillaire , 1824 (= Geocyon ; Pleistocene to this day in Africa)

Tribal history

Skull of the early hyena Ictitherium viverrinum in the American Museum of Natural History in New York

The hyenas first appeared in the Miocene of Europe around 17 million years ago and in the course of their tribal history gave birth to around 70 species, four of which are still alive today. The oldest genera are Protictitherium , Tungurictis and Plioviverrops from the Lower and Middle Miocene . These early hyenas were much smaller than today's species, about the size of a fox . Outwardly, Protictitherium looked more like a sneak cat and was probably an insect or omnivore that lived partially on trees. Plioviverrops , on the other hand, was more soil-oriented and resembled a mongoose . Both genera are likely to have had their origin in Western Europe and were common in Europe and Western Asia since the Middle Miocene. At the end of the Middle Miocene, Thalassictis , a somewhat larger genus, appeared in Europe. Thalassictis weighed about 20-30 kg and had dog-like teeth. Tungurictis , on the other hand, was widespread in East Asia at that time and had more pronounced hypercarnivore properties compared to its Western Eurasian relatives. Perhaps Tungurictis made from a Middle Miocene, emerged eastward expansion of early hyenas. The different adaptations to meat diet can also be an expression of deviating paleoecological conditions between western and eastern Eurasia. Accordingly, Tungurictis lived more in open landscapes, which is also indicated by his more developed toe gait.

The first hyenas were not specialized scavengers whose teeth were designed to bite into larger bones. Nevertheless, there were already large carnivores in the Middle Miocene, which looked similar to modern hyenas and probably occupied their ecological niche , but did not belong to the hyenas. Various hyena-like animals of the genera Percrocuta and Dinocrocuta , which were previously included in the hyenas, are now regarded as a separate family of predators, Percrocutidae . These predators resembled hyenas particularly in their skulls. The huge Dinocrocuta from the late Miocene , possibly weighing up to 380 kg , already had a highly developed scissor bite for cracking bones, as is known from various hyenas.

In the Upper Miocene, Ictitherium and Hyaenotherium, two other relatively large hyena genera, appeared in Europe. Both are likely to have originated in West Asia. Ictitherium was a little more omnivorous and had quite large molars. Hyaenotherium, on the other hand, was very much like Thalassictis , but was even larger. At the same time Protictitherium expanded its range and reached North Africa. Shortly afterwards, Adcrocuta, a really large genus of hyena appeared, which already had a specialized, bone-cracking bite. Adcrocuta reached a body weight of about 70 kg. The appearance of these large, bone-cracking hyenas may have initiated the decline of the Percrucotidae at the end of the Miocene. Another branch of hyenas produced relatively large, dog-like hyenas at the same time that were geared towards fast, persistent running. An early representative of this group was Hyaenictis . Another was Lycyaena .

Hyaenictis survived in Africa together with the dog-like form Hyaenictitherium into the Lower Pliocene , but disappeared shortly afterwards. Most of these dog-like hyenas were replaced at the transition from the Miocene to the Pliocene by representatives of the dogs (Canidae), who immigrated to the Old World from America for the first time at this time . While the hyenas developed in Eurasia and Africa in the Miocene and never reached the American continent with one exception, the dogs had developed in North America at the same time. The two families are very similar in their evolutionary history and each produced types that can be viewed as convergent . Among the early hyenas there were numerous forms that strongly resembled dogs, while in the Miocene and Pliocene of North America, instead of bone-cracking hyenas, scavenging dogs of the genus Osteoborus lived. In contrast to the hyenas adapted to running fast, which were largely replaced by dogs, the scavenging forms survived and produced larger and larger forms. These bones cracking hyenas, their precursors emerged since the Pliocene, was in addition to current species Crocuta and Hyaena also pachycrocuta . Pachycrocuta was the largest hyena of all time and weighed half more than today's spotted hyenas. The genus survived in Africa and Eurasia into the Pleistocene . Ikelohyaena abronia is considered to be a possible ancestor of today's striped hyenas and saddleback hyenas . A genus of forms adapted to walking, Chasmaporthetes , still occurred in the Pliocene and even reached America over the Bering Bridge . She was the only hyena that ever reached the New World . The northernmost known find of the genus is in the Old Crow Basin in the Yukon Territory in Canada, where some of the approximately 50,000 mammal finds that have been recovered also come from the hyena shape. It penetrated south into central Mexico. However, the appearance of Chasmaporthetes did not last very long in North America, as the genus there disappeared again in the course of the Middle Pleistocene . It is believed that it the pressure of competition from other large carnivores such as the giant short-faced bears could not withstand.

Aardwolves ( Proteles ) are known for the first time from the earliest Pleistocene of South Africa. Hyenas were also found in Europe in the late Pleistocene. The cave hyena , mostly regarded as a subspecies of the spotted hyena ( Crocuta crocuta ), was much larger than today's spotted hyenas and only died out at the end of the epoch.

literature

• Kay E. Holekamp, ​​Joseph M. Kolowski: Family Hyaenidae (Hyenas). In: Don E. Wilson, Russell A. Mittermeier (eds.): Carnivores (= Handbook of the Mammals of the World. Volume 1), Lynx Edicions, Barcelona 2009, ISBN 978-84-96553-49-1 , pp. 234-261.
• Markus Krajewski, Harun Maye (ed.): The hyena. Readings of a political animal . Diaphanes, Zurich 2010, ISBN 978-3-03734-136-0 .
• Ronald M. Nowak: Walker's Mammals of the World . Johns Hopkins University Press, Baltimore 1999, ISBN 0-8018-5789-9 .
• Jordi Agustí, Mauricio Antón: Mammoths, Sabertooths and Hominids. 65 Million Years of Mammalian Evolution in Europe. Columbia University Press, New York 2002, ISBN 0-231-11640-3 .

Web links

Commons : Hyaenidae  - Collection of images, videos and audio files

Wiktionary: Hyena  - explanations of meanings, word origins, synonyms, translations

• Information of Hyaena Specialist Group of the IUCN (English)
• Brown Hyena Research Project (English)

Individual evidence

1. ↑ Figures from Holekamp and Kolowski (2009).
2. ↑ ^{a b c} Andrea Naica-Loebell: Hyena women determine where to go. Heise, August 18, 2007
3. ^ Harald Schliemann: Carnivora, predators. In: Wilfried Westheide and Reinhard Rieger (eds.): Special Zoology Part 2: Vertebrate or Skull Animals , Gustav Fischer Verlag, Stuttgart Jena & New York 2003, ISBN 3-8274-0900-4 , p. 594.
4. ↑ Holekamp & Kolowski (2009), p. 245.
5. ↑ Holekamp & Kolowski (2009), p. 242.
6. ↑ Holekamp & Kolowski (2009), p. 259.
7. ↑ Holekamp & Kolowski (2009), p. 240.
8. ↑ Holekamp & Kolowski (2009), p. 254.
9. ^ W. Andrew Taylor and John D. Skinner: Associative feeding between Aardwolves (Proteles cristatus) and Aardvarks (Orycteropus afer). In: Mammal Review. 30 (2), 2000, pp. 141-143.
10. ↑ Holekamp and Kolowski (2009), p. 253.
11. ^ Günther Drosdowski (arrangement): Duden Etymologie. Dictionary of origin of the German language. 2nd Edition. Dudenverlag, Mannheim 2007, ISBN 3-411-20907-0 ( Der Duden. Volume 7).
12. ↑ Alma Gottlieb: Dog: Ally or Traitor? Mythology, Cosmology, and Society among the Beng of Ivory Coast. In: American Ethnologist. 13 (3), 1986, pp. 477-488, here p. 586.
13. ↑ Luc de Heusch: Sacrifice in Africa: A Structuralistic Approach. Manchester University Press, Manchester 1986, pp. 203f.
14. ^ ^{A b} Jürgen W. Frembgen : The Magicality of the Hyena: Beliefs and Practices in West and South Asia ( Memento from June 25, 2008 in the Internet Archive ). In: Asian Folklore Studies. 57, 1998, pp. 331-344, here p. 334.
15. ↑ Historia animalium VIII, 5; English online edition  ( page no longer available , search in web archives )  Info: The link was automatically marked as defective. Please check the link according to the instructions and then remove this notice. , accessed September 22, 2009.  
16. ^ ^{A b c} Stephen E. Glickman: The spotted hyena from Aristotle to the Lion King: Reputation is everything - In the Company of Animals. ( Memento from July 12, 2012 in the web archive archive.today ) In: Social Research. Fall 1995, accessed September 22, 2009.
17. ↑ The Millstätter Physiologus: Text, Translation, Commentary Edition on Google Books, pp. 82–85 , accessed on September 22, 2009.
18. ↑ The Letter to Barnabas. Greek-German text
19. ^ Mary Pendergraft: "Thou Shalt Not Eat the Hyena". A Note on “Barnabas” Epistle 10.7. In: Vigiliae Christianae. 46 (1), 1992, pp. 75-79.
20. ↑ * Hyenas in the Gutenberg-DE Brehms Thierleben project: online edition from the Gutenberg-DE project, accessed on September 23, 2009.
21. ↑ Holekamp & Kolowski (2009), p. 250.
22. ↑ Holekamp & Kolowski (2009), p. 257.
23. ^ Ingo Rieger: Hyaena hyaena. (PDF; 642 kB) In: Mammalian Species . No. 150, 1981, pp. 1-5.
24. ↑ ^{a b} Hyenas in the IUCN Red List of Endangered Species . Retrieved September 23, 2009.
25. ↑ Data of the spotted hyena according to Crocuta crocuta in the Red List of Threatened Species of the IUCN 2009. Posted by: O. Honer, KE Holekamp, ​​G. Mills, 2008. Retrieved on November 5, 2009., the other species according to Holekamp & Kolowski (2009 ), P. 251.
26. ↑ Yu Zhou, Si-Rui Wang, Jian-Zhang Ma: Comprehensive species set revealing the phylogeny and biogeography of Feliformia (Mammalia, Carnivora) based on mitochondrial DNA. In: PLoS ONE. 12 (3), 2017, p. E0174902 doi: 10.1371 / journal.pone.0174902 .
27. ↑ ^{a b c} Klaus-Peter Koepfli, Susan M. Jenks, Eduardo Eizirik, Tannaz Zahirpour, Blaire Van Valkenburgh, Robert K. Wayne: Molecular systematics of the Hyaenidae: Relationships of a relictual lineage resolved by a molecular supermatrix. In: Molecular Phylogenetics and Evolution. 38, 2006, pp. 603-620 and Anne D. Yoder, Melissa M. Burns, Sarah Zehr, Thomas Delefosse, Geraldine Veron, Steven M. Goodman, John J. Flynn: Single origin of Malagasy Carnivora from an African ancestor. (PDF; 388 kB) In: Nature . 421, 2003, pp. 734-737. Koepfli et al. a. examined only the fossa of the Malagasy carnivores that they believed to be a representative of the crawling cats; Yoder u. a. then all species of this group, but with the spotted hyena only one species of hyena.
28. ↑ e.g. Koehler & Richardson (1990)
29. ↑ for example: WC Wozencraft: Order Carnivora. In: DE Wilson, DM Reeder (Ed.): Mammal Species of the World . Smithsonian Institution Press, Washington 1993, pp. 279-344 or Holekamp & Kolowski (2009).
30. ↑ Lars Werdelin, Nikos Solounias: The Hyaenidae: taxonomy, systematlcs and evolution. In: Fossils and Strata. 30, 1991, pp. 1-104.
31. ↑ Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level in Columbia University Press, New York 1997, 631 pages, ISBN 0-231-11013-8 .
32. ↑ Don E. Wilson, DeeAnn M. Reeder (Ed.): Mammal Species of the World. A Taxonomic and Geographic Reference . 3. Edition. Johns Hopkins University Press, Baltimore 2005, ISBN 0-8018-8221-4 (2142 pages).
33. ↑ Lars Werdelin, Stéphane Peigné: Carnivora. In: Lars Werdelin, William Joseph Sanders (eds.): Cenozoic mammals of Africa. University of California Press, 2010, pp. 603-657.
34. ^ Lars Werdelin, Björn Kurtén: Allohyaena (Mammalia: Carnivora): giant hyaenid from the Late Miocene of Hungary. In: Zoological Journal of the Linnean Society. 126 (3), 1999, pp. 319-334.
35. ^ ^{A b c d} Alan Turner, Mauricio Antón: Evolving Eden. An Illustrated Guide to the Evolution of the African Large-Mammal Fauna. Columbia University Press, New York 2004, ISBN 0-231-11944-5 , pp. 78-81.
36. ^ ^{A b c} Xiaoming Wang, Richard H. Tedford, Mauricio Antón: Dogs. Their Fossil Relatives and Evolutionary History. Columbia University Press, New York 2008, ISBN 978-0-231-13528-3 , pp. 12-13.
37. ↑ Agustí (2002), p. 123.
38. ↑ Agustí (2002), p. 137.
39. ↑ Agustí (2002), p. 145.
40. ↑ Wang Xiaoming, Tseng Z. Jack, Wu Wen-Yu, Ye Jie, Meng Jin and Bi Shundong: A new species of Tungurictis Colbert, 1939 (Carnivora, Hyaenidae) from the middle Miocene of Junggar Basin, northwestern China and the early divergence of basal hyaenids in East Asia. In: Geodiversitas. 42 (3), 2020, pp. 29-45, doi: 10.5252 / geodiversitas2020v42a3
41. ↑ Agustí (2002), pp. 137-138.
42. ↑ Deng Tao, Tseng Zhijie J: Osteological evidence for predatory behavior of the giant percrocutid (Dinocrocuta gigantea) as an active hunter. In: Chinese Science Bulletin. 55 (17), 2010, pp. 1790-1794 (p. 1792).
43. ↑ Agustí (2002), p. 159.
44. ↑ Agustí (2002), pp. 174, 191, 193.
45. ↑ ^{a b} Agustí (2002), p. 213.
46. ^ Z. Jack Tseng, Grant Zazula, Lars Werdelin: First Fossils of Hyenas (Chasmaporthetes, Hyaenidae, Carnivora) from North of the Arctic Circle .. In: Open Quaternary. 5, 2019, p. 6, doi: 10.5334 / oq.64 .
47. ↑ Agustí (2002), p. 268.

This article was added to the list of excellent articles on November 30, 2009 in this version .