Equus gmelini
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The term tarpan (''Equus ferus ferus'') refers to free-ranging horses of the Russian steppe from the 18th to the 20th century. It is generally unknown whether those horses represented genuine wild horses, feral domestic horses or hybrids. The last individual believed to be a tarpan died in captivity in the Russian Empire during 1909. Beginning in the 1930s, several attempts were made to develop horses that looked like tarpans through selective breeding, called "
breeding back Breeding back is a form of artificial selection by the deliberate selective breeding of domestic (but not exclusively) animals, in an attempt to achieve an animal breed with a phenotype that resembles a wild type ancestor, usually one that has g ...
" by advocates. The breeds that resulted included the
Heck horse The Heck horse is a horse breed that is claimed to resemble the tarpan (''Equus ferus ferus''), an extinct equine. The breed was created by the German zoologist brothers Heinz Heck and Lutz Heck in an attempt to breed back the tarpan. Althoug ...
, the , 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 Kazakh, Qazaq or Kazakhstani may refer to: * Someone or something related to Kazakhstan *Kazakhs, an ethnic group *Kazakh language *The Kazakh Khanate * Kazakh cuisine * Qazakh Rayon, Azerbaijan *Qazax, Azerbaijan *Kazakh Uyezd, administrative dis ...
or
Kyrgyz Kyrgyz, Kirghiz or Kyrgyzstani may refer to: * Someone or something related to Kyrgyzstan *Kyrgyz people *Kyrgyz national games *Kyrgyz language *Kyrgyz culture *Kyrgyz cuisine *Yenisei Kirghiz *The Fuyü Gïrgïs language in Northeastern China ...
) name meaning " wild horse". The Tatars and the
Cossacks The Cossacks , es, cosaco , et, Kasakad, cazacii , fi, Kasakat, cazacii , french: cosaques , hu, kozákok, cazacii , it, cosacchi , orv, коза́ки, pl, Kozacy , pt, cossacos , ro, cazaci , russian: казаки́ or ...
distinguished the wild horse from the feral horse; the latter was called ''Takja'' or ''Muzin''.


Taxonomy

The tarpan was first described by
Samuel Gottlieb Gmelin Samuel George Gottlieb Gmelin (4 July 1744 – 27 July 1774) was a German physician, botanist, and explorer. Background Gmelin was born at Tübingen as part of a well-known family of naturalists. His father was Johann Conrad Gmelin, an apothecar ...
in 1771; 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 Otto Antonius (21 May 1885 in Vienna – 9 April 1945 in Vienna) was director of the Tiergarten Schönbrunn in Vienna, zoologist, palaeontologist and co-founder of the modern zoological biology. Early life Otto Antonius was the eldest of fi ...
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. 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. 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. In 2003, the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
"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.


Appearance

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''. The last individual, which died in captivity in 1909, was between 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. A number of coat color
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
s have been identified within European wild horses from the Pleistocene and Holocene: those creating
bay A bay is a recessed, coastal body of water that directly connects to a larger main body of water, such as an ocean, a lake, or another bay. A large bay is usually called a Gulf (geography), gulf, sea, sound (geography), sound, or bight (geogra ...
, black and leopard complex are known from the wild horse population in Europe and were depicted in
cave painting In archaeology, Cave paintings are a type of parietal art (which category also includes petroglyphs, or engravings), found on the wall or ceilings of caves. The term usually implies prehistoric origin, and the oldest known are more than 40,000 ye ...
s of wild horses during the Pleistocene. The dun gene, a
dilution gene A dilution gene is any one of a number of genes that act to create a lighter coat color in living creatures. There are many examples of such genes: General Diluted coat colors have melanocytes, but vary from darker colors due to the concentration ...
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. Whether or not the horses called tarpan had a standing mane like wild equines, or a falling mane like domestic horses cannot be ascertained anymore as historic reports have not been unambiguous on this subject. The appearance of European wild horses may be reconstructed with genetic, osteologic and historic data. One genetic study suggests that bay was the predominant color in European wild horses. During the
Mesolithic The Mesolithic (Greek: μέσος, ''mesos'' 'middle' + λίθος, ''lithos'' 'stone') or Middle Stone Age is the Old World archaeological period between the Upper Paleolithic and the Neolithic. The term Epipaleolithic is often used synonymous ...
, a gene coding a black coat color appeared on the Iberian peninsula. This color spread east but was less common than bay in the investigated sample and never reached Siberia. Bay in combination with dun results in the "bay dun" color 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 colors were a better camouflage in forests.Baker, Sue, 2008: Exmoor Ponies: Survival of the Fittest – A natural history. Pangaré or "mealy" coloration, a characteristic of other wild equines, might have been present in at least some European wild horses, as historic accounts report a light belly. Historic references describing the tarpan report that most of them were of a black dun (grullo, "blue dun") colour, while also black individuals and domestic colours like white legs or greys were reported. Because of that, some contemporaneous authors, such as Peter Pallas, believed tarpans to be escaped farm horses.Tadeusz Jezierski, Zbigniew Jaworski: Das Polnische Konik. Die Neue Brehm-Bücherei Bd. 658, Westarp Wissenschaften, Hohenwarsleben 2008


History

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 Russia, where a horse of this type was domesticated around 3000 BC. ''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.


Forest tarpan

The "forest horse" or "forest tarpan" was a hypothesis of various 20th-century natural scientists, including
Tadeusz Vetulani Tadeusz Bolesław Vetulani (13 March 1897 – 24 February 1952) was a Polish agriculturalist and biologist, associate professor of Adam Mickiewicz University in animal husbandry. He was a pioneer of biodiversity research in Poland and conducted not ...
, 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.'' 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 The Ardennes (french: Ardenne ; nl, Ardennen ; german: Ardennen; wa, Årdene ; lb, Ardennen ), also known as the Ardennes Forest or Forest of Ardennes, is a region of extensive forests, rough terrain, rolling hills and ridges primarily in Be ...
, Great Britain, and the southern
Swedish Swedish or ' may refer to: Anything from or related to Sweden, a country in Northern Europe. Or, specifically: * Swedish language, a North Germanic language spoken primarily in Sweden and Finland ** Swedish alphabet, the official alphabet used by ...
upland. They had a robust head and strong body, and a long frizzy mane. The color was described as faint brown or yellowish brown with
eel stripe 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 ...
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. Yet, those horses were never colloquially called "tarpans".Cis van Vuure: On the origin of the Polish konik and its relation to Dutch nature management. 2014 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. 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. 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 Samuel George Gottlieb Gmelin (4 July 1744 – 27 July 1774) was a German physician, botanist, and explorer. Background Gmelin was born at Tübingen as part of a well-known family of naturalists. His father was Johann Conrad Gmelin, an apothecar ...
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-colored with a light belly and legs becoming black, although gray and white horses were mentioned as well. The coat was long and dense. The horses at Zamosc were never called "tarpan" back in their lifetime. 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-colored legs or gray coats.


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. They often were mixed with domestic horses, and alongside pure herds there were herds of feral horses or hybrids. The color of alleged pure tarpans was described as consistently brown, cream-colored or mouse-colored. 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. According to Smith, tarpans made stronger sounds than domestic horses and the overall appearance of these horses was mule-like.


Extinction

The human-caused extinction of wild horses in Europe began in Southern Europe, possibly in antiquity. While humans had been hunting wild horses since the
Paleolithic The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός ''palaios'', "old" and λίθος ''lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone too ...
, 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
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 of such matings were intractable. Tarpans lived in the southern parts of the Russian steppe. In 1879 the last scientifically confirmed individual was killed. After that, only dubious sightings were documented. 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 Russian zoo. The horses at Zamosc survived until 1806, when they were allegedly donated to local farmers of Poland and bred to their horses.Bunzel-Drüke, Finck, Kämmer, Luick, Reisinger, Riecken, Riedl, Scharf & Zimball: ''"Wilde Weiden: Praxisleitfaden für Ganzjahresbeweidung in Naturschutz und Landschaftsentwicklung''Thomas Jansen: ''Untersuchungen zur Phylogenie und Domestikation des Hauspferdes (Equus ferus f. caballus) Stammesentwicklung und geografische Verteilung.'' 2002
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The Konik is claimed to descend from these hybrid horses, as recent research has highlighted a significant degree of anatomic difference between free-roaming Konik in the Netherlands and other modern domesticated horses.


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). The diverse
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 ...
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, and local use of wild mares or even secondary sites of domestication are likely. Therefore, the European wild horse may well have contributed to the domestic horse.


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. Historic references to "wild horses" may actually refer to feral domestic horses or hybrids. Some 19th century authors wrote that local "wild" horses had hoof problems that led to crippled legs; therefore, they assumed these were feral horses. 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. 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. The only known individual to be photographed was the so-called Cherson tarpan, which was caught as a foal near
Novovorontsovka Novovorontsovka ( uk, Нововоронцовка; russian: Нововоронцовка) is an urban-type settlement in Beryslav Raion, Kherson Oblast, southern Ukraine. It hosts the administration of Novovorontsovka settlement hromada, one of t ...
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. 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, 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.


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 Tadeusz Bolesław Vetulani (13 March 1897 – 24 February 1952) was a Polish agriculturalist and biologist, associate professor of Adam Mickiewicz University in animal husbandry. He was a pioneer of biodiversity research in Poland and conducted not ...
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,Tadeusz Jezierski, Zbigniew Jaworski: ''Das Polnische Konik. Die Neue Brehm-Bücherei Bd. 658'', Westarp Wissenschaften, Hohenwarsleben 2008, which clusters genetically with other domestic horse breeds, including those as diverse as the Mongolian horse and the Thoroughbred. In the early 1930s,
Berlin Zoo The Berlin Zoological Garden (german: link=no, Zoologischer Garten Berlin) is the oldest surviving and best-known zoo in Germany. Opened in 1844, it covers and is located in Berlin's Tiergarten. With about 1,380 different species and over 20,2 ...
Director
Lutz Heck Ludwig Georg Heinrich Heck, called Lutz Heck (23 April 1892 in Berlin, German Empire – 6 April 1983 in Wiesbaden, West Germany) was a German zoologist, animal researcher, animal book author and director of the Berlin Zoological Garden where he ...
and
Heinz Heck Heinz Heck (22 January 1894 – 5 March 1982) was a German biologist and director of Hellabrunn Zoo (''Tierpark Hellabrunn'') in Munich. He was born in Berlin and died in Munich. With his brother, Lutz Heck, who was director of the Berlin ...
of the
Munich Zoo Hellabrunn Zoo (or Tierpark Hellabrunn in German) is a 40 hectare (99 acre) zoological garden in the Bavarian capital of Munich. The zoo is situated on the right bank of the river Isar, in the southern part of Munich near the quarter of Thalkirc ...
began a program crossbreeding Koniks with Przewalski horses, Gotland ponies, and Icelandic horses. By the 1960s they produced the
Heck horse The Heck horse is a horse breed that is claimed to resemble the tarpan (''Equus ferus ferus''), an extinct equine. The breed was created by the German zoologist brothers Heinz Heck and Lutz Heck in an attempt to breed back the tarpan. Althoug ...
. In the mid-1960s, Harry Hegard started a similar program in the United States using
mustangs The mustang is a free-roaming horse of the Western United States, descended from horses brought to the Americas by the Spanish. Mustangs are often referred to as wild horses, but because they are descended from once-domesticated animals, they ...
and local working ranch horses that has resulted in the Hegardt or Stroebel's horse.


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". In spite of sharing primitive external features, the Konik and
Hucul The Hutsuls (sometimes the spelling variant: Gutsuls; uk, Гуцули, translit=Hutsuly; pl, Huculi, Hucułowie; ro, huțuli) are an ethnic group spanning parts of western Ukraine and Romania (i.e. parts of Bukovina and Maramureș). They ...
horses have markedly different conformation with differently proportioned body measurements, thought in part to be linked to living in different habitats. Other breeds sometimes alleged to be surviving wild horses include the
Exmoor pony The Exmoor pony is one of the mountain and moorland pony breeds native to the British Isles. Some still roam as semi-feral livestock on Exmoor, a large area of moorland in Devon and Somerset in southwest England. The Exmoor has been given "P ...
and the
Dülmen pony Dülmen () is a town in the district of Coesfeld, North Rhine-Westphalia, Germany. Geography Dülmen is situated in the south part of the Münsterland area, between the Lippe river to the south, the Baumberge hills to the north and the Ems riv ...
. However, genetic studies do not set any of these breeds apart from other domestic horses. On the other hand, there has not yet been a study comparing domestic breeds directly with the European wild horse.


See also

* Domestication of the horse * List of extinct animals of Europe * Przewalski's horse


References


External links


The Spanish TarpanThe Spanish Tarpan in North AmericaAmerican "Tarpan"North American Tarpan Association
{{short description, Extinct subspecies of equines Controversial mammal taxa Equus (genus) Extinct mammals of Europe Species made extinct by human activities Mammal extinctions since 1500 Pleistocene horses Horse subspecies Mammals described in 1785