Tarpan

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Tarpan
Temporal range: Holocene
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Kherson tarpan.jpg
The "Cherson tarpan", the only tarpan to be photographed, 1884
Status iucn3.1 EX.svg
Extinct  (1909)  (IUCN 3.1)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Equidae
Genus: Equus
Species:
E. ferus
Subspecies:
†E. f. ferus
Trinomial name
†Equus ferus ferus
Boddaert, 1785
Synonyms

The tarpan (Equus ferus ferus) was a free-ranging horse population of the Eurasian steppe from the 18th to the 20th century. [1] What qualifies as a tarpan is subject to confusion. It is unknown whether those horses represented genuine wild horses, feral domestic horses or hybrids. [1] [2] The last individual believed to be a tarpan died in captivity in the Russian Empire in 1909. [3]

Contents

Beginning in the 1930s, several attempts were made to develop horses that looked like tarpans through selective breeding, called "breeding back" by advocates. The breeds that resulted included the Heck horse, the Hegardt or Stroebel's horse  [ fr ], and a derivation of the Konik breed, all of which have a primitive appearance, particularly in having the grullo coat colour. Some of these horses are now commercially promoted as "tarpans", although such animals are only domestic breeds and not the wild animal themselves.

Name and etymology

The name "tarpan" or "tarpani" derives from a Turkic language (Kazakh or Kyrgyz) name meaning "wild horse". [4] [5] The Tatars and the Cossacks distinguished the wild horse from the feral horse; the latter was called Takja or Muzin. [6] [7]

Taxonomy

The tarpan was first described by Samuel Gottlieb Gmelin in 1771; [8] he had seen the animals in 1769 in the district of Bobrov, near Voronezh. In 1784, Pieter Boddaert named the species Equus ferus, referring to Gmelin's description. Unaware of Boddaert's name, Otto Antonius published the name Equus gmelini in 1912, again referring to Gmelin's description. Since Antonius' name refers to the same description as Boddaert's it is a junior objective synonym. It is now thought that the domesticated horse, named Equus caballus by Carl Linnaeus in 1758, is descended from the tarpan; indeed, many taxonomists consider them to belong to the same species. By a strict application of the rules of the International Code of Zoological Nomenclature, the tarpan ought to be named E. caballus, or if considered a subspecies, E. caballus ferus. However, biologists have generally ignored the letter of the rule and used E. ferus for the tarpan to avoid confusion with the domesticated subspecies.[ citation needed ]

It is debated if the small, free-roaming horses seen in the Russian steppes during 18th and 19th centuries and called "tarpan" were indeed wild, never-domesticated horses, hybrids of the Przewalski's horse and local domestic animals, or simply feral horses. [9] Most studies have been based on only two preserved specimens and research to date has not positively linked the tarpan to Pleistocene or Holocene-era animals.[ citation needed ]

In 2003, the International Commission on Zoological Nomenclature "conserved the usage of 17 specific names based on wild species, which are predated by or contemporary with those based on domestic forms", confirming E. ferus for the undomesticated wild horse. Taxonomists who consider the domestic horse a subspecies of the wild horse should use Equus ferus caballus; the name Equus caballus remains available for the domestic horse where it is considered to be a separate species. [10]

Appearance

Only known illustration of a tarpan made from life, depicting a five-month-old foal. By Borisov, 1841 Tarpan.png
Only known illustration of a tarpan made from life, depicting a five-month-old foal. By Borisov, 1841

Traditionally, two tarpan subtypes have been proposed, the forest tarpan and steppe tarpan, although there seem to be only minor differences in type. The general view is that there was only one subspecies, the tarpan, Equus ferus ferus. [3] The last individual, which died in captivity in 1909, was between 140 and 145 centimetres (55 and 57 in; 13.3 and 14.1 hands ) tall at the shoulders, and had a thick, falling mane, a grullo coat colour, dark legs, and primitive markings, including a dorsal stripe and shoulder stripes. [3]

A number of coat colour genotypes have been identified within European wild horses from the Pleistocene and Holocene: those creating bay, black and leopard complex are known from the wild horse population in Europe and were depicted in cave paintings of wild horses during the Pleistocene. [12] The dun gene, [13] a dilution gene seen in Przewalski's horse that also creates the grullo or "blue dun" coat seen in the Konik, has not yet been studied in European wild horses. It is likely that at least some wild horses had a dun coat. [12]

Historical reports are ambiguous on whether tarpans had standing manes like wild equines, or falling manes like domestic horses. [3]

The appearance of European wild horses may be reconstructed with genetic, osteologic and historic data. One genetic study suggests that bay was the predominant colour in European wild horses. [14] During the Mesolithic, a gene coding a black coat colour appeared on the Iberian peninsula. This colour spread east but was less common than bay in the investigated sample and never reached Siberia. [14] Bay in combination with dun results in the "bay dun" colour seen in Przewalski's horses; black with dun creates the grullo coat. A loss of the dun dilution may have been advantageous in more forested western European landscapes, as dark colours were a better camouflage in forests. [15] Pangaré or "mealy" coloration, a characteristic of other wild equines, might have been present in at least some European wild horses, [15] as historic accounts report a light belly.

Historic references report that most tarpans were black dun (grullo, "blue dun") colour. Some black individuals were reported to have domestic colours like white or grey legs. Authors, such as Peter Pallas, believed tarpans to be escaped farm horses. [16]

History

Replica of a horse painting from a cave in Lascaux Lascaux horse replica.jpg
Replica of a horse painting from a cave in Lascaux

Wild horses have been present in Europe since the Pleistocene and ranged from southern France and Spain east to central Russia. There are cave drawings of primitive predomestication horses at Lascaux, France and in Cave of Altamira, Spain, as well as artifacts believed to show the species in southern Russian empire, where a horse of this type was domesticated around 3000 BC. [17] Equus ferus had a continuous range from western Europe to Alaska; historic material suggests wild horses lived in most parts of Holocene continental Europe and the Eurasian steppe, except for parts of Scandinavia, Iceland and Ireland. [16]

DNA

DNA obtained from a "tarpan" that lived in the steppes of the Kherson region during the mid-19th century shows its ancestry to be a mixture between horses native to Europe and found with the Corded Ware Culture, and horses closely related to the DOM2 population, the ancestors of modern domestic horses. (DOM2 horses originated from the Western Eurasia steppes, especially the lower Volga-Don, but not in Anatolia, during the late fourth and early third millennia BCE. Their genes may show selection for easier domestication and stronger backs). This is not consistent with tarpans as the wild ancestor of, or a feral version of, DOM2. Nor is the tarpan a hybrid with Przewalski's horses. [18]

Forest tarpan

The "forest horse" or "forest tarpan" was a hypothesis of various 20th-century natural scientists, including Tadeusz Vetulani, who suggested that the continuous forestation of Europe after the last ice age created forest-adapted subtype of the wild horse, which he named Equus sylvestris. However, historic references do not describe any major difference between the populations, and therefore most authors assume there was only one subspecies of western Eurasian wild horse, Equus ferus ferus. [3]

Nevertheless, a stocky type of horse living in forests and highlands was described during the 19th century in Spain, the Pyrenees, the Camargue, the Ardennes, Great Britain, and the southern Swedish upland. They had a robust head and strong body, and a long frizzy mane. The colour was described as faint brown or yellowish brown with eel stripe and leg stripes, or wholly black legs. The flanks and shoulders were spotted, some of them tended to an ashy colour. They dwelled in rocky habitats and showed intelligent and fierce behaviour. [7] Yet, those horses were never colloquially called "tarpans". [19]

Black wild horses were found in Dutch swamps, with a large skull, small eyes, and a bristly muzzle. Their mane was full, with broad hooves, and curly hair. However, it is possible that these were feral and not wild horses. [7]

European wild horse coat colours Wild horse reconstruction.jpg
European wild horse coat colours

Herodotus described light-coloured wild horses in an area now part of Ukraine in the 5th century BCE. In the 12th century, Albertus Magnus stated that mouse-coloured wild horses with a dark eel stripe lived in the German territory, and in Denmark, large herds were hunted.

16th century free-ranging horses in Europe

Wild horses still were common in the east of Prussia during the 15th and early 16th centuries. During the 16th century, wild horses disappeared from most of the mainland of western Europe and became less common in eastern Europe as well. Belsazar Hacquet saw wild horses in the Polish zoo in Zamość during the Seven Years' War. According to him, those wild horses were of small body size, had a blackish brown colour, a large and thick head, short dark manes and tail hair, and a "beard". They were absolutely untameable and defended themselves harshly against predators.[ citation needed ]

Kajetan Kozmian visited the population at Zamość as well and reported that they were small and strong, had robust limbs and a consistently dark mouse colour. Samuel Gottlieb Gmelin witnessed herds in Voronezh in 1768. Those horses were described as very fast and shy, fleeing at any noise, and as small, with small, pinned ears and a short frizzly mane. The tail was shorter than in domestic horses. They were typically mouse-coloured with a light belly and legs becoming black, although grey and white horses were mentioned as well. The coat was long and dense. [16] The horses at Zamosc were never called "tarpan" back in their lifetime. [19]

Peter Simon Pallas witnessed possible tarpans in the same year in southern Russia. He thought they were feral animals that escaped during the confusions of wars. These herds were important game of the Tatars and numbered between 5 and 20 animals. The horses he described had a small body, large and thick heads, short frizzly manes and short tail hair, as well as pinned ears. The colour was described as faint brownish, sometimes brown or black. He also reported of obvious domestic hybrids with light-coloured legs or grey coats. [16]

The tarpans of 18th century Europe

The Natural History of Horses by 19th-century author Charles Hamilton Smith also described tarpans. According to Smith, the herds of free-ranging horses numbered from a few to hundred individuals. [7] They often were mixed with domestic horses, and alongside pure herds there were herds of feral horses or hybrids. The colour of alleged pure tarpans was described as consistently brown, cream-coloured or mouse-coloured. The short frizzy mane was reported to be black, as were the tail and legs. The ears were of varying size, but set high on the skull. The eyes were small. [16]

According to Smith, tarpans made stronger sounds than domestic horses and the overall appearance of these horses was mule-like. [7]

Extinction

Illustration of a running individual Equus ferus ferus.jpg
Illustration of a running individual

The human-caused extinction of wild horses in Europe began in Southern Europe, possibly in antiquity. [16] While humans had been hunting wild horses since the Paleolithic, [15] during historic times horse meat was an important source of protein for many cultures. As large herbivores, the range of the tarpan was continuously decreased by the increasing human population of the Eurasian landmass. Wild horses were further targeted because they caused damage to hay stores and often took domestic mares from pastures. Furthermore, interbreeding with wild horses was an economic loss for farmers since the foals of such matings were intractable.[ further explanation needed ] [16]

Tarpans lived in the southern parts of the steppe. In 1879 the last scientifically confirmed individual was killed. After that, only dubious sightings were documented. [16] As the tarpan horse died out in the wild between 1875 and 1890, the last free-ranging mare considered a tarpan was accidentally killed during an attempt at capture. The last captive tarpan died in 1909 in a Ukrainian zoo in the Russian empire. [21]

The horses at Zamosc survived until 1806, when they were allegedly donated to local farmers of Poland and bred to their horses. [3] [22] The Konik is claimed to descend from these hybrid horses, [17] as recent research has highlighted a significant degree of anatomic difference between free-roaming Konik in the Netherlands and other modern domesticated horses. [23]

Tarpans interbreeding with domestic horses

The oldest archaeological evidence for domesticated horses is from Kazakhstan and Ukraine between 6,000 and 5,500  YBP (years before present). [24] The diverse mitochondrial DNA of domestic horses contrasts sharply with the very low diversity of the Y chromosome; that suggests that many mares but only a few stallions were used, [25] and local use of wild mares or even secondary sites of domestication are likely. [26] Therefore, the European wild horse may well have contributed to the domestic horse. [26]

Wild horses vs. feral horses

Some researchers consider the tarpans to be mixed wild and feral population or completely feral horses. Few consider the more recent animals historically called "tarpans" to be genuine wild horses without domestic influence. [16] Historic references to "wild horses" may actually refer to feral domestic horses or hybrids. [16]

Some 19th century authors wrote that local "wild" horses had hoof problems that led to crippled legs; therefore, they assumed these were feral horses. [16] Other contemporary authors claimed all "wild" horses between the Volga River and the Ural were actually feral. However, others thought that this was too speculative and assumed that wild, undomesticated horses still lived into the 19th century. [7] Domestic horses used in warfare often were turned loose when they were no longer needed. Also, remaining wild stallions could steal domestic mares. There are some accounts from the 18th and 19th centuries of wild herds with typical wild horse features such as large heads, pinned ears, short frizzy mane and tail, but mentioned animals with domestic influence as well. [16]

The only known individual to be photographed was the so-called Cherson tarpan, which was caught as a foal near Novovorontsovka in 1866. It died in 1887 in the Moscow Zoo. The nature of this horse was dubious in its lifetime, because it showed almost none of the wild horse features described in the historic sources. [3] In 2021, a study found that the so-called 'Shatilov' tarpan, a museum specimen from the Kherson region of the Pontic–Caspian steppe that died in 1868[ citation needed ], was a hybrid between two horse lineages, with two thirds of its genetics representing the same ancestral lineage as modern horses, and the remaining third related to a distinct lineage of European horses found at an archaeological site associated with the late-Neolithic Corded Ware culture. [27]

Heck horse in Haselunne, Germany (2004) Steppentarpan.jpg
Heck horse in Haselünne, Germany (2004)

Breeding back the tarpan

Three attempts have been made to use selective breeding to create a type of horse that resembles the tarpan phenotype, though recreating an extinct subspecies is not genetically possible with current technology. In 1936, Polish university professor Tadeusz Vetulani selected Polish farm horses that were formerly known as Panje horses (now called Konik) and that he believed resembled the historic tarpan and started a selective breeding program. Although his breeding attempt is well known, it made only a minor contribution to the modern Konik stock, [1] which clusters genetically with other domestic horse breeds, including those as diverse as the Mongolian horse and the Thoroughbred. [26]

In the early 1930s, Berlin Zoo Director Lutz Heck and Heinz Heck of the Munich Zoo began a program crossbreeding Koniks with Przewalski horse stallions, and the mares of Gotland ponies, and Icelandic horses,. By the 1960s they produced the Heck horse. [28] In the mid-1960s, Harry Hegard started a similar program in the United States using mustangs and local working ranch horses that has resulted in the Hegardt or Stroebel's horse. [29]

Assessment

While all three breeds have a primitive look that resembles the tarpan in some respects, they are not genetically tarpans and the wild, predomestic European horse remains extinct. However, this does not prevent some modern breeders from marketing horses with these features as a "tarpan". [30]

In spite of sharing primitive external features, the Konik and Hucul horses have markedly different conformation with differently proportioned body measurements, thought in part to be linked to living in different habitats. [3] [31]

Other breeds sometimes alleged to be surviving wild horses include the Exmoor pony and the Dülmen pony. However, genetic studies do not set any of these breeds apart from other domestic horses. [26] [32] On the other hand, there has not yet been a study comparing domestic breeds directly with the European wild horse. [33]

See also

Related Research Articles

<span class="mw-page-title-main">Horse</span> Domesticated equine

The horse is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of Equus ferus. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature, Eohippus, into the large, single-toed animal of today. Humans began domesticating horses around 4000 BCE, and their domestication is believed to have been widespread by 3000 BCE. Horses in the subspecies caballus are domesticated, although some domesticated populations live in the wild as feral horses. These feral populations are not true wild horses, which are horses that never have been domesticated. There is an extensive, specialized vocabulary used to describe equine-related concepts, covering everything from anatomy to life stages, size, colors, markings, breeds, locomotion, and behavior.

<span class="mw-page-title-main">Domestication of the horse</span>

How and when horses became domesticated has been disputed. Although horses appeared in Paleolithic cave art as early as 30,000 BC, these were wild horses and were probably hunted for meat. The clearest evidence of early use of the horse as a means of transport is from chariot burials dated c. 2000 BC. However, an increasing amount of evidence began to support the hypothesis that horses were domesticated in the Eurasian Steppes in approximately 3500 BC. Discoveries in the context of the Botai culture had suggested that Botai settlements in the Akmola Province of Kazakhstan are the location of the earliest domestication of the horse. Warmouth et al. (2012) pointed to horses having been domesticated around 3000 BC in what is now Ukraine and Western Kazakhstan.

<span class="mw-page-title-main">Przewalski's horse</span> Subspecies of mammal

Przewalski's horse, also called the takhi, Mongolian wild horse or Dzungarian horse, is a rare and endangered horse originally native to the steppes of Central Asia. It is named after the Russian geographer and explorer Nikolay Przhevalsky. Once extinct in the wild, since the 1990s it has been reintroduced to its native habitat in Mongolia in the Khustain Nuruu National Park, Takhin Tal Nature Reserve, and Khomiin Tal, as well as several other locales in Central Asia and Eastern Europe.

<span class="mw-page-title-main">Breeding back</span> Artificial selection

Breeding back is a form of artificial selection by the deliberate selective breeding of domestic animals, in an attempt to achieve an animal breed with a phenotype that resembles a wild type ancestor, usually one that has gone extinct. Breeding back is not to be confused with dedomestication.

<span class="mw-page-title-main">Konik</span> Polish breed of horse

The Konik or Polish Konik, Polish: konik polski, is a Polish breed of small horse or pony. There are semi-feral populations in some regions. They are usually mouse dun or striped dun.

<i>Equus</i> (genus) Genus of mammals which includes horses, donkeys, and zebras

Equus is a genus of mammals in the family Equidae, which includes horses, asses, and zebras. Within the Equidae, Equus is the only recognized extant genus, comprising seven living species. Like Equidae more broadly, Equus has numerous extinct species known only from fossils. The genus originated in North America and dispersed into the Old World and South America during the Early and Middle Pleistocene. Equines are odd-toed ungulates with slender legs, long heads, relatively long necks, manes, and long tails. All species are herbivorous, and mostly grazers, with simpler digestive systems than ruminants but able to subsist on lower-quality vegetation.

<span class="mw-page-title-main">Equine coat color genetics</span> Genetics behind the equine coat color

Equine coat color genetics determine a horse's coat color. Many colors are possible, but all variations are produced by changes in only a few genes. Bay is the most common color of horse, followed by black and chestnut. A change at the agouti locus is capable of turning bay to black, while a mutation at the extension locus can turn bay or black to chestnut.

<span class="mw-page-title-main">Dun gene</span> Dilution gene

The dun gene is a dilution gene that affects both red and black pigments in the coat color of a horse. The dun gene lightens most of the body while leaving the mane, tail, legs, and primitive markings the shade of the undiluted base coat color. A dun horse always has a dark dorsal stripe down the middle of its back, usually has a darker face and legs, and may have transverse striping across the shoulders or horizontal striping on the back of the forelegs. Body color depends on the underlying coat color genetics. A classic "bay dun" is a gray-gold or tan, characterized by a body color ranging from sandy yellow to reddish brown. Duns with a chestnut base may appear a light tan shade, and those with black base coloration are a smoky gray. Manes, tails, primitive markings, and other dark areas are usually the shade of the undiluted base coat color. The dun gene may interact with all other coat color alleles.

<span class="mw-page-title-main">Sorraia</span> Breed of horse

The Sorraia is a rare breed of horse indigenous to the portion of the Iberian Peninsula, in the Sorraia River basin, in Portugal. The Sorraia is known for its primitive features, including a convex profile and dun coloring with primitive markings. Concerning its origins, a theory has been advanced by some authors that the Sorraia is a descendant of primitive horses belonging to the naturally occurring wild fauna of Southern Iberia. Studies are currently ongoing to discover the relationship between the Sorraia and various wild horse types, as well as its relationship with other breeds from the Iberian Peninsula and Northern Africa.

<span class="mw-page-title-main">Wild horse</span> Undomesticated four-footed mammal from the equine family

The wild horse is a species of the genus Equus, which includes as subspecies the modern domesticated horse as well as the endangered Przewalski's horse. The European wild horse, also known as the tarpan, that went extinct in the late 19th or early 20th century has previously been treated as the nominate subspecies of wild horse, Equus ferus ferus, but more recent studies have cast doubt on whether tarpans were truly wild or if they actually were feral horses or hybrids.

<span class="mw-page-title-main">Grullo</span> Color of horses in the dun family

Grullo or grulla is a color of horses in the dun family, characterized by tan-gray or mouse-colored hairs on the body, often with shoulder and dorsal stripes and black barring on the lower legs. The genotype for grulla horses is a black base with dun dilution. In this coloration, each individual hair is mouse-colored, unlike a roan, which is composed of a mixture of dark and light hairs. The several shades of grulla are informally referred to with a variety of terms, including black dun, blue dun, slate grulla, silver grulla or light grulla, silver dun, or lobo dun. Silver grulla may also refer to a grulla horse with silver dapple, regardless of shade.

<i>Equus lambei</i> Extinct species of horse

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<span class="mw-page-title-main">Mongolian horse</span> Breed of horse

The Mongolian horse is the native horse breed of Mongolia. The breed is purported to be largely unchanged since the time of Genghis Khan. Nomads living in the traditional Mongol fashion still hold more than 3 million animals, which outnumber the country's human population. In Mongolia, the horses live outdoors all year, dealing with temperatures from 30 °C (86 °F) in summer down to −40 °C (−40 °F) in winter, and they graze and search for food on their own. The mare's milk is processed into the national beverage airag. Some animals are slaughtered for meat. Other than that, they serve as riding and transport animals; they are used both for the daily work of the nomads and in horse racing.

<span class="mw-page-title-main">Feral horse</span> Free horses of domesticated horse ancestry

A feral horse is a free-roaming horse of domesticated stock. As such, a feral horse is not a wild animal in the sense of an animal without domesticated ancestors. However, some populations of feral horses are managed as wildlife, and these horses often are popularly called "wild" horses. Feral horses are descended from domestic horses that strayed, escaped, or were deliberately released into the wild and remained to survive and reproduce there. Away from humans, over time, these animals' patterns of behavior revert to behavior more closely resembling that of wild horses. Some horses that live in a feral state but may be occasionally handled or managed by humans, particularly if privately owned, are referred to as "semi-feral".

<span class="mw-page-title-main">Evolution of the horse</span>

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<span class="mw-page-title-main">Primitive markings</span> Hair coloration in some equine

Primitive markings are a group of hair coat markings and qualities seen in several equine species, including horses, donkeys, and asses. In horses, they are associated with primitive breeds, though not limited to such breeds. The markings are particularly associated with the dun coat color family. All dun horses possess at least the dorsal stripe, but the presence of the other primitive markings varies. Other common markings may include horizontal striping on the legs, transverse striping across the shoulders, and lighter guard hairs along the edges of a dark mane and tail.

<span class="mw-page-title-main">Heck horse</span> Horse breed

The Heck horse is a horse breed that is claimed to resemble the tarpan, an extinct wild equine. The breed was created by the German zoologist brothers Heinz Heck and Lutz Heck in an attempt to breed back the tarpan. Although unsuccessful at creating a genetic copy of the extinct species, they developed a look-alike breed with grullo coloration and primitive markings. Heck horses were subsequently exported to the United States, where a breed association was created in the 1960s.

Tarpan is an extinct Eurasian wild horse.

<span class="mw-page-title-main">Pony</span> Type of small horse

A pony is a type of small horse. Depending on the context, a pony may be a horse that is under a given height at the withers, or a small horse with a specific conformation and temperament. Compared to a larger horse, a pony may have a thicker coat, mane and tail, with proportionally shorter legs, a wider barrel, heavier bone, a thicker neck and a shorter, broader head. The word pony derives from the old French poulenet, meaning foal, a young, immature horse.

The history of horse domestication has been subject to much debate, with various competing hypotheses over time about how domestication of the horse occurred. The main point of contention was whether the domestication of the horse occurred once in a single domestication event, or that the horse was domesticated independently multiple times. The debate was resolved at the beginning of the 21st century using DNA evidence that favored a mixed model in which domestication of the stallion most likely occurred only once, while wild mares of various regions were included in local domesticated herds.

References

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  2. ↑ Lovász, Lilla; Fages, Antoine; Amrhein, Valentin (December 2021). "Konik, Tarpan, European wild horse: An origin story with conservation implications". Global Ecology and Conservation. 32: e01911. doi:10.1016/j.gecco.2021.e01911.
  3. 1 2 3 4 5 6 7 8 Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: "Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung
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  5. ↑ "Tarpan". Vasmer's Etymological Dictionary.
  6. ↑ Boyd, Lee; Houpt, Katherine A. (1994). Przewalski's Horse: The History and Biology of an Endangered Species. SUNY Series in Endangered Species. Albany State University of New York Press. ISBN   0-7914-1890-1.
  7. 1 2 3 4 5 6 Smith, Charles Hamilton (1866) [1841]. The Natural History of Horses, with Memoir of Gesner.
  8. ↑ "Comparative craniometry of "Shatilov's tarpan" (Equus gmelini Antonius, 1912): a problem of species status" (PDF). Zoological Museum of Moscow University. 49. 2008. Archived from the original (PDF) on 2019-11-10. Retrieved 2020-03-26.
  9. ↑ Castelli, Andrea (2016). "Don't Call Me Tarpan". The Official PLOS Blog (published May 24, 2016).
  10. ↑ International Commission on Zoological Nomenclature (2003). Opinion 2027 (Case 3010). "Usage of 17 specific names based on wild species which are predated by or contemporary with those based on domestic animals (Lepidoptera, Osteichthyes, Mammalia): conserved." Bulletin of Zoologic Nomenclature, 60:81–84.
  11. ↑ Bennett, 1998
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  22. ↑ Thomas Jansen: Untersuchungen zur Phylogenie und Domestikation des Hauspferdes (Equus ferus f. caballus) Stammesentwicklung und geografische Verteilung. 2002 (PDF Archived 2016-03-08 at the Wayback Machine )
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