Equines of Africa
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''Equus'' , is a genus of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s in the family Equidae, which includes horses, donkeys, and zebras. Within the Equidae, ''Equus'' is the only recognized
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
genus, comprising seven living species. Like Equidae more broadly, ''Equus'' has numerous
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
species known only from fossils. The genus most likely originated in North America and spread quickly to the Old World. Equines are odd-toed ungulates with slender legs, long heads, relatively long necks, manes (erect in most subspecies), and long tails. All species are herbivorous, and mostly grazers, with simpler digestive systems than ruminants but able to subsist on lower-quality vegetation. While the domestic horse and donkey (along with their feral descendants) exist worldwide, wild equine populations are limited to Africa and Asia. Wild equine social systems are in two forms; a harem system with tight-knit groups consisting of one adult male or
stallion A stallion is a male horse that has not been gelded (castrated). Stallions follow the conformation and phenotype of their breed, but within that standard, the presence of hormones such as testosterone may give stallions a thicker, "cresty" nec ...
, several females or mares, and their young or
foal A foal is an equine up to one year old; this term is used mainly for horses, but can be used for donkeys. More specific terms are colt for a male foal and filly for a female foal, and are used until the horse is three or four. When the foal i ...
s; and a territorial system where males establish territories with resources that attract females, which associate very fluidly. In both systems, females take care of their offspring, but males may play a role as well. Equines communicate with each other both visually and vocally. Human activities have threatened wild equine populations.


Etymology

The word is Latin for "horse" and is cognate with the Greek (, "horse") and Mycenaean Greek , the earliest attested variant of the Greek word, written in
Linear B Linear B was a syllabic script used for writing in Mycenaean Greek, the earliest attested form of Greek. The script predates the Greek alphabet by several centuries. The oldest Mycenaean writing dates to about 1400 BC. It is descended from ...
syllabic script. Compare the alternative development of the Proto-Greek labiovelar in Ionic ().


Taxonomic and evolutionary history

The genus ''Equus'' was first described by Carl Linnaeus in
1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of Systema Naturae, 10th edition of ''Systema Naturae'', the sta ...
. It is the only recognized
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
genus in the family Equidae. The first equids were small, dog-sized mammals (e.g. '' Eohippus'') adapted for browsing on shrubs during the Eocene, around 54 million years ago (Mya). These animals had three toes on the hind feet and four on the front feet with small hooves in place of claws, but also had soft pads. Equids developed into larger, three-toed animals (e.g. '' Mesohippus'') during the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
and Miocene. From there, the side toes became progressively smaller through the Pleistocene until the emergence of the single-toed ''Equus''. The genus ''Equus'', which includes all extant equines, is believed to have evolved from ''
Dinohippus ''Dinohippus'' (Greek: "Terrible horse") is an extinct equid which was endemic to North America from the late Hemphillian stage of the Miocene through the Zanclean stage of the Pliocene (10.3—3.6 mya) and in existence for approximately . Fossi ...
'', via the intermediate form ''
Plesippus ''Plesippus'' is a genus of extinct horse from the Pleistocene of North America. Although commonly seen as a subgenus of ''Equus'' recent cladistic analysis considers it a distinct genus. Species Two species are recognized by Barron et al. (201 ...
''. One of the oldest species is '' Equus simplicidens'', described as zebra-like with a donkey-like head shape. The oldest material to date was found in Idaho, USA. The genus appears to have spread quickly into the Old World, with the similarly aged ''E. livenzovensis'' documented from western Europe and Russia. Molecular phylogenies indicate that the most recent common ancestor of all modern equines (members of the genus ''Equus'') lived ~5.6 (3.9-7.8) Mya. Direct paleogenomic sequencing of a 700,000-year-old middle Pleistocene horse metapodial bone from Canada implies a more recent 4.07 Mya for the most recent common ancestor within the range of 4.0 to 4.5 Mya. Mitochondrial evidence supports the division of ''Equus'' species into noncaballoid (which includes zebras and asses) and caballoids or "true horses" (which includes '' E. ferus'' and '' E. przewalskii''). Of the extant equine species, the lineage of the asses may have diverged first, possibly as soon as ''Equus'' reached the Old World. Zebras appear to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
and differentiated in Africa, where they are endemic. Members of the subgenus ''Sussemionus'' were abundant during the Early and Middle Pleistocene of North America and Afro-Eurasia, but only a single species, ''E. ovodovi'' survived into the Late Pleistocene and Holocene in south Siberia and China, with the youngest remains from China dating to around 3500 BP (1500 BC), during the Shang dynasty. Genetic data from ''E. ovodovi'' has placed the ''Sussemionus'' lineage as closer to zebras and asses than to caballine horses. Molecular dating indicates the caballoid lineage diverged from the noncaballoids 4 Mya. Genetic results suggest that all North American fossils of caballine equines, as well as South American fossils traditionally placed in the subgenus ''E.'' ('' Amerhippus''), belong to ''E. ferus''. Remains attributed to a variety of species and lumped together as
New World stilt-legged horse ''Haringtonhippus'' is an extinct genus of stilt-legged equine from the Pleistocene of North America The genus is monospecific, consisting of the species ''H. francisci'', initially described in 1915 by Oliver Perry Hay as ''Equus francisci''. ...
s (including ''E. francisci'', ''E. tau'', and ''E. quinni'') probably all belong to a second species that was endemic to North America. This was confirmed in a genetic study done in 2017, which subsumed all the specimens into the species ''E. francisci'' which was placed outside all extant horse species in the new genus '' Haringtonhippus','' although its placement as a separate genus was subsequently questioned. A separate genus of horse, '' Hippidion'' existed in South America. The possible causes of the extinction of horses in the Americas (about 12,000 years ago) have been a matter of debate. Hypotheses include
climatic change ''Climatic Change'' is a biweekly peer-reviewed scientific journal published by Springer Science+Business Media covering cross-disciplinary work on all aspects of climate change and variability. It was established in 1978 and the editors-in-chief ...
and
overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...
by newly arrived humans. Horses only returned to the American mainland with the arrival of the ''
conquistador Conquistadors (, ) or conquistadores (, ; meaning 'conquerors') were the explorer-soldiers of the Spanish and Portuguese Empires of the 15th and 16th centuries. During the Age of Discovery, conquistadors sailed beyond Europe to the Americas, O ...
es'' in 1519.


Extant species


Prehistoric Species

Many extinct prehistoric species of ''Equus'' have been described. The validity of some of these species is questionable and a matter of debate. For example, ''Equus niobrarensis'' is likely synonymous with ''Equus scotti'', while ''Equus alaskae'' is most likely the same species as ''Equus lambei'', which itself may be a North American form of the living ''Equus przewalskii''. DNA studies on American horse remains found frozen into
permafrost Permafrost is ground that continuously remains below 0 °C (32 °F) for two or more years, located on land or under the ocean. Most common in the Northern Hemisphere, around 15% of the Northern Hemisphere or 11% of the global surface ...
have shown that several of the supposed American species, and the European '' Equus ferus'', are actually one highly-variable widespread species. , as if the evolutionary process of
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
was persistently being frustrated by large herds of the horses moving long distances and mixing, carrying their
genes In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
about with them. *''
Equus alaskae ''Equus alaskae'' was a Pleistocene species of horse, now extinct, that inhabited North America. Fossils found from Alaska to Mexico have been identified as ''Equus alaskae,'' and it has been referred to as the most common equid in the southwest ...
'' - Alaskan horse *''
Equus algericus Equus may refer to: * ''Equus'' (genus), a genus of animals including horses, donkeys and zebras * ''Equus'' (play), a play by Peter Shaffer * ''Equus'' (film), a film adaptation of the Peter Shaffer play * Equus (comics), a comic book characte ...
'' *''
Equus capensis ''Equus capensis'' (''E. capensis''), the 'giant Cape zebra', is an extinct species of zebra that lived during the Pleistocene of South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmo ...
'' - Giant zebra *''
Equus conversidens ''Equus conversidens'', or the Mexican horse, is a dubious Pleistocene species of horse, now extinct, that inhabited North America. The holotype of ''Equus conversidens'', a partial palate, was unearthed in Pleistocene deposits northeast of Mexi ...
'' - Mexican horse *'' Equus dalianensis'' *''
Equus lenensis ''Equus lenensis'', the Lena horse, is an extinct species of horse from the Late Pleistocene and Holocene of Siberia, Some sources have considered it a subspecies of the wild horse. Genetic studies show that ''E. lenensis'' does not descend from ...
'' - Siberian horse *''
Equus latipes Equus may refer to: * ''Equus'' (genus), a genus of animals including horses, donkeys and zebras * ''Equus'' (play), a play by Peter Shaffer * ''Equus'' (film), a film adaptation of the Peter Shaffer play * Equus (comics), a comic book characte ...
'' *''
Equus fraternus ''Equus fraternus'' is an extinct species of '' Equus,'' which was native to North America. Specimens of ''E. fraternus'' have been found in Florida, Louisiana and Virginia Virginia, officially the Commonwealth of Virginia, is a state in ...
'' *''
Equus giganteus The giant horse (''Equus giganteus'') is an extinct species of horse which lived in North America. It was classified as a species based on the finding of a single tooth larger than the teeth of even the largest modern draft horse A draft h ...
'' - Giant horse *''
Equus lambei ''Equus lambei'', commonly known as the Yukon horse or Yukon wild horse, is an extinct species of the genus '' Equus''. ''Equus lambei'' ranged across North America until approximately 10,000 years ago. Based on recent examinations of the m ...
'' - Yukon horse *''
Equus mauritanicus Equus may refer to: * ''Equus'' (genus), a genus of animals including horses, donkeys and zebras * ''Equus'' (play), a play by Peter Shaffer * ''Equus'' (film), a film adaptation of the Peter Shaffer play * Equus (comics), a comic book characte ...
'' - Saharan zebra *''
Equus namadicus ''Equus namadicus'' is a prehistoric equid, known from remains dating to the Middle and Late Pleistocene from across the Indian subcontinent The Indian subcontinent is a physiographical region in Southern Asia. It is situated on the I ...
'' *''
Equus neogeus ''Amerhippus'' is an extinct South American horse of uncertain taxonomic identity. It is sometimes classified as a subgenus of the genus '' Equus'', containing several extinct species of horses that lived in South America, or a single, morphologi ...
'' - often placed in a separate genus, '' Amerhippus'' *''
Equus niobrarensis ''Equus niobrarensis'' (or commonly, Niobrara horse) is an extinct species of '' Equus'', the genus that includes the horse. ''E. niobrarensis'' may be synonymous with ''Equus scotti''. It was "stout-legged" and belonged to the "big horses" cat ...
'' - Niobrara horse *'' Equus occidentalis'' - Western horse *''
Equus ovodovi Equus may refer to: * Equus (genus), ''Equus'' (genus), a genus of animals including horses, donkeys and zebras * Equus (play), ''Equus'' (play), a play by Peter Shaffer * Equus (film), ''Equus'' (film), a film adaptation of the Peter Shaffer play ...
'' - Ovodov's horse *'' Equus scotti'' - Scott's horse *''
Equus semiplicatus ''Equus semplicatus'', was a Pleistocene species of New World stilt-legged horse, and considered the type species for the stilt legged horses, one of three lineages of equids within the Americas, the other two being hippidionid and caballine ho ...
'' *'' Equus simplicidens'' - Hagerman horse *''
Equus sivalensis ''Equus sivalensis'' is an extinct equid, discovered in the Siwalik hills. Remains date to 2.6 million years ago, and it is assumed that it was extinct during the last ice age, between 75,000 and 10,000 years ago, as part of the late Pleistoce ...
'' - Indian horse *''
Equus stenonis ''Equus stenonis'' or the Stenon zebra, is an extinct species of equine closely related to modern zebras and asses that inhabited Eurasia in the Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch ...
'' - Stenon zebra *''
Equus yunnanensis The Yunnan horse (''Equus yunnanensis'') was an extinct horse that once roamed in Pleistocene East AsiaElewa, Ashraf M. T. (2008). Mass Extinction. Springer p. 172Colbert, EH. (1940)Pleistocene mammals from the Ma Kai valley of northern Yunnan, ...
'' - Yunnan horse


Hybrids

Equine species can crossbreed with each other. The most common
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...
is the
mule The mule is a domestic equine hybrid between a donkey and a horse. It is the offspring of a male donkey (a jack) and a female horse (a mare). The horse and the donkey are different species, with different numbers of chromosomes; of the two pos ...
, a cross between a male donkey and a female horse. With rare exceptions, these hybrids are
sterile Sterile or sterility may refer to: *Asepsis, a state of being free from biological contaminants * Sterile (archaeology), a sediment deposit which contains no evidence of human activity *Sterilization (microbiology), any process that eliminates or ...
and cannot reproduce. A related hybrid, a hinny, is a cross between a male horse and a female donkey. Other hybrids include the
zorse A zebroid is the offspring of any cross between a zebra and any other equine to create a hybrid. In most cases, the sire is a zebra stallion. Offspring of a donkey sire and zebra dam are called a donkra and offspring of a horse sire and a zebra ...
, a cross between a zebra and a horse and a zonkey or zedonk, a hybrid of a zebra and a donkey. In areas where Grévy's zebras are
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
with plains zebras, fertile hybrids do occur. Ancient DNA identifies the Bronze Age kunga as a cross between the Syrian wild ass and the donkey.


Biology


Physical characteristics

Equines have significant differences in size, though all are characterized by long heads and necks. Their slender legs support their weight on one digit (which evolved from the middle digits). Grévy's zebra is the largest wild species, standing up to and weighing up to . Domesticated horses have a wider range of sizes. Heavy or draft horses are usually at least high and can be as tall as and weigh from about . Some miniature horses are no taller than in adulthood. Sexual dimorphism is limited in equines. The penis of the male is vascular and lacks a bone (
baculum The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the ...
). Equines are adapted for running and traveling over long distances. Their
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
is adapted for grazing; they have large incisors that clip grass blades and highly crowned, ridged molars well suited for grinding. Males have spade-shaped canines ("tushes"), which can be used as weapons in fighting. Equines have fairly good senses, particularly their eyesight. Their moderately long, erect ears are movable and can locate the source of a sound. A dun-colored coat with primitive markings that include a dorsal stripe and often leg striping and transverse shoulder stripes reflect the wildtype coat and are observed in most wild extant equine species. Only the mountain zebra lacks a dorsal stripe. In domestic horses, dun color and primitive markings exist in some animals across many breeds. The purpose of the bold black-and-white striping of zebras has been a subject of debate among biologists for over a century, but 2014 evidence supports the theory that they are a form of protection from biting flies. These insects appear to be less attracted to striped coats, and compared to other wild equines, zebras live in areas with the highest fly activity. With the exception of the domestic horses, which have long manes that lay over the neck and long tail hair growing from the top of the tailhead or
dock A dock (from Dutch language, Dutch ''dok'') is the area of water between or next to one or a group of human-made structures that are involved in the handling of boats or ships (usually on or near a shore) or such structures themselves. The ex ...
, most equines have erect manes and long tails ending in a tuft of hair. The coats of some equine species undergo shedding in certain parts of their range and are thick in the winter.


Ecology and daily activities

Extant wild equines have scattered ranges across Africa and Asia. The plains zebra lives in lush grasslands and savannas of Eastern and Southern Africa, while the mountain zebra inhabits mountainous areas of southwest Africa. The other equine species tend to occupy more arid environments with more scattered vegetation. Grévy's zebra is found in thorny scrubland of East Africa, while the African wild ass inhabits rocky deserts of North Africa. The two Asian wild ass species live in the dry deserts of the Near East and Central Asia and Przwelski's wild horse's habitat is the deserts of Mongolia. Only the range of the plains and Grévy's zebras overlap. In addition to wild populations, domesticated horses and donkeys are widespread due to humans. In certain parts of the world, populations of feral horses and feral donkeys exist, which are descended from domesticated animals that were released or escaped into the wild. Equines are monogastric
hindgut fermenters Hindgut fermentation is a digestive process seen in monogastric herbivores, animals with a simple, single-chambered stomach. Cellulose is digested with the aid of symbiotic bacteria.ruminants, equines have a simpler and less efficient digestive system. Nevertheless, they can subsist on lower-quality vegetation. After food is passed through the stomach, it enters the sac-like cecum, where cellulose is broken down by micro-organisms. Fermentation is quicker in equines than in ruminants—30–45 hours for a horse compared to 70–100 hours for cattle. Equines may spend 60–80% of their time feeding, depending on the availability and quality of vegetation. In the African savannas, the plains zebra is a pioneer grazer, mowing down the upper, less nutritious grass canopy and preparing the way for more specialized grazers such as blue wildebeests and Thomson's gazelles, which depend on shorter and more nutritious grasses below. Wild equines may spend seven hours a day sleeping. During the day, they sleep standing up, while at night they lie down. They regularly rub against trees, rocks, and other objects and roll in around in dust for protection against flies and irritation. Except the mountain zebra, wild equines can roll over completely.


Social behavior

Equines are social animals with two basic social structures. Horses, plains zebras, and mountain zebras live in stable, closed family groups or harems consisting of one adult male, several females, and their offspring. These groups have their own home ranges, which overlap and they tend to be nomadic. The stability of the group remains even when the family stallion dies or is displaced. Plains zebra groups gather into large herds and may create temporarily stable subgroups within a herd, allowing individuals to interact with those outside their group. Among harem-holding species, this behavior has only otherwise been observed in primates such as the gelada and the
hamadryas baboon The hamadryas baboon (''Papio hamadryas'' ) is a species of baboon within the Old World monkey family. It is the northernmost of all the baboons, being native to the Horn of Africa and the southwestern region of the Arabian Peninsula. These re ...
. Females of harem species benefit as males give them more time for feeding, protection for their young, and protection from predators and harassment by outside males. Among females in a harem, a linear dominance hierarchy exists based on the time at which they join the group. Harems travel in a consistent filing order with the high-ranking mares and their offspring leading the groups followed by the next-highest ranking mare and her offspring, and so on. The family stallion takes up the rear.
Social grooming Social grooming is a behavior in which social animals, including humans, clean or maintain one another's body or appearance. A related term, allogrooming, indicates social grooming between members of the same species. Grooming is a major soci ...
(which involves individuals rubbing their heads against each other and nipping with the incisors and lips) is important for easing aggression and maintaining social bonds and status. Young of both sexes leave their natal groups as they mature; females are usually abducted by outside males to be included as permanent members of their harems. In Grévy's zebras and the wild ass species, adults have more fluid associations and adult males establish large
territories A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or an ...
and monopolize the females that enter them. These species live in habitats with sparser resources and standing water, and grazing areas may be separated. Groups of lactating females are able to remain in groups with nonlactating ones and usually gather at foraging areas. The most dominant males establish territories near watering holes, where more sexually receptive females gather. Subdominants have territories farther away, near foraging areas. Mares may wander through several territories, but remain in one when they have young. Staying in a territory offers a female protection from harassment by outside males, as well as access to a renewable resource. Some feral populations of horses exhibit features of both the harem and territorial social systems. In both equine social systems, excess males gather in bachelor groups. These are typically young males that are not yet ready to establish a harem or territory. With the plains zebra, the males in a bachelor group have strong bonds and have a linear dominance hierarchy. Fights between males usually occur over estrous females and involve biting and kicking.


Communication

When meeting for the first time or after they have separated, individuals may greet each other by rubbing and sniffing their noses followed by rubbing their cheeks, moving their noses along their bodies and sniffing each other's genitals. They then may rub and press their shoulders against each other and rest their heads on one another. This greeting is usually performed among harem or territorial males or among bachelor males playing. Equines produce a number of vocalizations and noises. Loud snorting is associated with alarm. Squealing is usually made when in pain, but bachelors also squeal while play fighting. The contact calls of equines vary from the whinnying and nickering of the horse and the barking of plains zebras to the braying of asses, Grévy's zebras, and donkeys. Equines also communicate with visual displays, and the flexibility of their lips allows them to make complex facial expressions. Visual displays also incorporate the positions of the head, ears, and tail. An equine may signal an intention to kick by laying back its ears and sometimes lashing the tail. Flattened ears, bared teeth, and abrupt movement of the heads may be used as threatening gestures, particularly among stallions.


Reproduction and parenting

Among harem-holding species, the adult females mate only with their harem stallion, while in other species, mating is more promiscuous and the males have larger testes for sperm competition. Estrus in female equines lasts 5–10 days; physical signs include frequent urination, flowing muscus, and swollen, everted labia. In addition, estrous females will stand with their hind legs spread and raise their tails when in the presence of a male. Males assess the female's reproductive state with the flehmen response and the female will solicit mating by backing in. Length of gestation varies by species; it is roughly 11–13 months, and most mares come into estrus again within a few days after foaling, depending on conditions. Usually, only a single foal is born, which is capable of running within an hour. Within a few weeks, foals attempt to graze, but may continue to nurse for 8–13 months. Species in arid habitats, like Grévy's zebra, have longer nursing intervals and do not drink water until they are three months old. Among harem-holding species, foals are cared for mostly by their mothers, but if threatened by predators, the entire group works together to protect all the young. The group forms a protective front with the foals in the center and the stallion will rush at predators that come too close. In territory-holding species, mothers may gather into small groups and leave their young in " kindergartens" under the guard of a territorial male while searching for water. Grévy's zebra stallions may look after a foal in his territory to ensure that the mother stays, though it may not be his.


Human relations

The earliest archaeological evidence for the domestication of the horse comes from sites in Ukraine and Kazakhstan, dating to around 4000–3500 BCE. By 3000 BCE, the horse was completely domesticated, and by 2000 BCE, a sharp increase occurred in the number of horse bones found in human settlements in northwestern Europe, indicating the spread of domesticated horses throughout the continent. The most recent, but most irrefutable, evidence of domestication comes from sites where horse remains were buried with chariots in graves of the Sintashta and Petrovka cultures c. 2100 BCE. Studies of variation in genetic material shows that a very few wild stallions, possibly all from a single
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 or ...
, contributed to the domestic horse, mating with many mares in early domesticated herds. Przewalski's horse has been conclusively shown not to be an ancestor of the domestic horse, though the two can hybridize and produce fertile offspring. The split between Przewalskii's horse and ''E. ferus caballus'' is estimated to have occurred 120,000–240,000 years ago, long before domestication. Of the caballine equines of ''E. ferus,'' ''E. f. ferus,'' also known as the European wild horse or "tarpan", shares ancestry with the modern domestic horse. In addition, tarpans that lived into modern times may have been hybridized with domestic horses. Archaeological, biogeographical, and linguistic evidence suggests that the donkey was first domesticated by nomadic pastoral people in North Africa over 5,000 years ago. The animals were used to help cope with the increased aridity of the Sahara and the
Horn of Africa The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004), ...
. Genetic evidence finds that the donkey was domesticated twice based on two distinct
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
haplogroups A haplotype is a group of alleles in an organism that are inherited together from a single parent, and a haplogroup ( haploid from the el, ἁπλοῦς, ''haploûs'', "onefold, simple" and en, group) is a group of similar haplotypes that shar ...
. It also points to a single ancestor, the Nubian wild ass. Attempts to domesticate zebras were largely unsuccessful, though
Walter Rothschild Lionel Walter Rothschild, 2nd Baron Rothschild, Baron de Rothschild, (8 February 1868 – 27 August 1937) was a British banker, politician, zoologist and soldier, who was a member of the Rothschild family. As a Zionist leader, he was present ...
trained some to draw a carriage in England.


Conservation issues

Humans have had a great impact on the populations of wild equines. Threats to wild equines include habitat destruction and conflicts with local people and livestock. Since the 20th century, wild equines have been decimated over many of their former ranges and their populations scattered. In recent centuries, two subspecies, the quagga and the tarpan, became extinct. 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 ...
lists the African wild ass as critically endangered, Grévy's zebra, the mountain zebra, and Przewalski's horse as endangered, the onager as
vulnerable Vulnerable may refer to: General * Vulnerability * Vulnerability (computing) * Vulnerable adult * Vulnerable species Music Albums * ''Vulnerable'' (Marvin Gaye album), 1997 * ''Vulnerable'' (Tricky album), 2003 * ''Vulnerable'' (The Used album) ...
, the plains zebra as near threatened, and the kiang as least concern. Przewalski's horse was considered to be
extinct in the wild A species that is extinct in the wild (EW) is one that has been categorized by the International Union for Conservation of Nature as known only by living members kept in captivity or as a naturalized population outside its historic range due ...
from the 1960s to 1996. However, following successful captive breeding, it has been reintroduced in Mongolia. Feral horses vary in degree of protection and generate considerable controversy. For example, in Australia, they are considered a non-native
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
, often viewed as pests, though are also considered to have some cultural and economic value. In the United States, feral horses and burros are generally considered an introduced species because they are descendants from domestic horses brought to the Americas from Europe. While they are viewed as pests by many livestock producers, conversely, a view also exists that ''E. f. caballus'' is a
reintroduced Species reintroduction is the deliberate release of a species into the wild, from captivity or other areas where the organism is capable of survival. The goal of species reintroduction is to establish a healthy, genetically diverse, self-sustainin ...
once-native species returned to the Americas that should be granted
endangered species An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
protection. At present, certain free-roaming horses and burros have federal protection as "living symbols of the historic and pioneer spirit of the West" under the Wild and Free-Roaming Horses and Burros Act of 1971, and in ''
Kleppe v. New Mexico ''Kleppe v. New Mexico'', 426 U.S. 529 (1976), was a United States Supreme Court decision that unanimously held the Wild and Free-Roaming Horses and Burros Act of 1971, passed in 1971 by the United States Congress to protect these animals from "ca ...
'', the United States Supreme Court ruled that the animals so designated were, as a matter of law, wildlife.


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{{DEFAULTSORT:Equus (Genus) Mammal genera Extant Calabrian first appearances Taxa named by Carl Linnaeus