Overview

Brief Summary

The tarpan or Eurasian wild horse, Equus caballus ferus or E. ferus, is an extinct wild horse related to the common domestic horse (E. caballus caballus) that roamed from southern France and Spain to central Russia, in steppe and forest environments.  They stood approximately 140-145 centimetres (55-57 inches) tall at the shoulders, had a partly falling mane, a mouse-grey (grullo) coat color, dark legs and primitive markings, including a dorsal stripe and shoulder stripes.  Smith (1866) described tarpans as mule-like, and making stronger sounds than domestic horses (Mass 2000-2013; Wikipedia 2014).

The name “tarpan” is Turkic for “wild horse,” and is distinguished from the feral horse (called Takja or Muzin).  In modern use, the term has been loosely used to refer to the pre-domesticated ancestor of the modern horse, Equus ferus; to the pre-domestic subspecies believed to have lived into the historic era, Equus caballus ferus; and nonspecifically to all European primitive or "wild" horses in general (Wikipedia 2014).  Other common synonyms for tarpans are Equus ferus ferus and E. c. gmelini.

Beginning in the 1930s, several attempts were made to genetically reconstruct horses that looked like tarpans through selective breeding, called “breeding back” by advocates.  The breeds that resulted include the Heck horse, the Hegardt or Stroebel's horse, and a derivation of the Konik breed, all of which have a primitive appearance, particularly in having the grullo coat color.  Some of these horses are now commercially promoted as "tarpans," although researchers discourage this use of the word, which they believe should only applied to the ancient E. caballus ferus (Castelli 2012).

Tarpans became extinct starting in Southern Europe, as a result of human hunting and a range decreasing in size with the increasing civilization of the Eurasian continent.  They were persecuted because they caused damage to hay storages, often took domestic mares from pastures and because interbreeding with wild horses was an economic loss for farmers since the foals of such matings were intractable.   Tarpans survived the longest in the southern parts of the Russian steppe.  By 1880, when most “Tarpans” may have become hybrids, wild horses were very rare.  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 considered-wild mare was accidentally killed during an attempt at capture.  The last captive individual believed to be a tarpan died in 1909 in a Russian zoo (Wikipedia 2014).

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Distribution

Range Description

Until the late 18th century, this species ranged from Germany and Russian Steppes east to Kazakhstan, Mongolia and northern China. After this time, the species went into catastrophic decline. Wild animals survived in eastern Europe (Poland, Belarus, Lithuania and Germany) through the eighteenth century, with the last wild individuals possibly killed in 1814 (Novak 1999). The Plains Tarpan (Equus ferus ferus), lived on the steppes of southern Russia and the Ukraine. The last wild population of Przewalski’s Horse survived until recently in southwestern Mongolia and adjacent Gansu, Xinjiang, and Inner Mongolia (China). Wild horses were last seen in 1969, north of the Tachiin Shaar Nuruu in Dzungarian Gobi Desert in Mongolia (Paklina and Pozdnyakova 1989).

Since the 1990s, reintroduction efforts have started in Mongolia, China, Kazakhstan and Ukraine; Mongolia is the only country where truly wild reintroduced populations exist within its historic range. Reintroductions in Mongolia began in Takhin Tal Nature Reserve in the Dzungarian Gobi Desert (9,000 km²) and Hustai National Park in Mongol Daguur Steppe (570 km²) in 1994 (King and Gurnell 2005). A third reintroduction site, Khomiin Tal, (2,500 km²), in the Great Lakes Depression, was established in 2004, as a buffer zone to the Khar Us Nuur National Park in Valley of the Lakes (C. Feh pers. comm.).

All living wild horses belong to the subspecies Equus ferus przewalskii. The first visual account of Przewalski’s-type wild horses date from more than 20,000 years ago. Rock engravings, paintings, and decorated tools dating from the late Gravetian to the late Magdalenian (20,000–9,000 BC), were discovered in caves in Italy, southern France, and northern Spain; 610 of these were horse figures (Leroi-Gourhan 1971). Many cave drawings in France show horses that look like Przewalski’s Horse (Mohr 1971). In prehistoric times, the species probably roamed widely over the steppes of central Asia, China, and Europe (Ryder 1990). The first written accounts originate from Tibet, recorded by the monk Bodowa, who lived around 900 AD. In the “Secret History of the Mongols”, there is also a reference to wild horses that crossed the path of Chinggis Khaan during his campaign against Tangut in 1226, causing his horse to rear and throw him to the ground (Bokonyi 1974). That the wild horse was a prestigious gift, denoting its rarity or that it was difficult to catch, is shown by the presentation of a Przewalski’s horse to the emperor of Manchuria by Chechen-Khansoloj-Chalkaskyden, an important Mongolian, circa 1630 (Zevegmid and Dawaa 1973). In a Manchurian dictionary of 1771, Przewalski’s Horse is mentioned as “a wild horse from the steppe” (Dovchin 1961).

Przewalski’s Horse was not described in Linnaeus’s “Systema Naturae” (1758) and remained largely unknown in the West until first mentioned by John Bell, a Scottish doctor who travelled in the service of Tsar Peter the Great in 1719–1722 (Mohr 1971). His account of the expedition, “A Journey from St Petersburg to Peking”, was published in 1763. Bell and subsequent observers all located horses known at that time within the area of 85–97°E and 43–50°N (Chinese-Mongolian border). Wild horses were reported again from what is now China by Colonel Nikolai Mikailovich Przewalski, an eminent explorer, at the end of the nineteenth century. He made several expeditions by order of Tsar Alexander the Second of Russia to central Asia, aiming to reach Tibet. While returning from his second expedition in central Asia, he was presented with the skull and hide of a horse shot about 80 km north of Gutschen on the Chinese-Russian border. The remains were examined at the Zoological Museum of the Academy of Science in St Petersburg by I.S. Poliakov, who concluded that they were a wild horse, which he gave the official name Equus przewalskii (Poliakov 1881). Further reports came from the brothers Grigory and Michael Grum-Grzhimailo, who travelled through western China from 1889-1890. In 1889, they discovered a group in the Gashun area and shot four horses: three stallions, and a mare. The four hides and the skulls of the three stallions, together with an incomplete skeleton, were sent back to the Zoological Museum in St. Petersburg. They were able to observe the horses from a short distance and gave the following account: “Wild horses keep in bands of no more than ten, each herd having a dominant stallion. There are other males, too, but they are young and, judging by the hide of the two-year old colt that we killed, the dominant male treats them very cruelly. In fact, the hide showed traces of numerous bites” (Grum-Grzhimailo 1982). Current scientific review of the taxonomy of wild equids (Groves 1986) places Przewalski’s Horse as a subspecies of Equus ferus.
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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology

Przewalski’s Horse formerly inhabited steppe and semi-desert habitats, as most of this range became degraded or was occupied by livestock, the species became restricted to semi-desert habitats with limited water resources (Van Dierendonck and de Vries 1996). Lowland steppe vegetation was preferentially selected by horses at Hustai National Park and seasonal movements are affected by the availability of the most nutritious vegetation (King and Gurnell 2005).

Because the historic range is not precisely known, there has been much debate about the areas in which Przewalski’s Horses were last seen: was it merely a last refuge or was it representative of the typical/preferred habitat? The Mongolia Takhi Strategy and Plan Work Group (MTSPWG 1993) concluded that the historic range may have been wider but that the Dzungarian Gobi, where they were last seen, was not a marginal site to which the species retreated. Although grass and water are more available in other parts of Mongolia, these areas often have much harsher winters

An alternative viewpoint of the desert-steppe controversy is that the Eurasian steppe should be considered the wild horse's optimal habitat (Van Dierendonck and de Vries 1996). This would suggest that Przewalski’s Horses were forced into sub-optimal ranges such as the arid Gobi, as the more favourable steppe region was colonized by nomadic pastoralist people over several millennia. Studies of feral horses have shown that they are able to live and reproduce in semi-desert habitats but their survival and reproductive success is clearly sub-optimal compared to feral horses on more mesic grassland (Berger 1986). Van Dierendonck and de Vries (1996) suggest that the wild horse is primarily a steppe herbivore that can survive under arid conditions when there is access to waterholes.

Systems
  • Terrestrial
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
EN
Endangered

Red List Criteria
D

Version
3.1

Year Assessed
2011

Assessor/s
Boyd, L. & King, S.R.B.

Reviewer/s
Moehlman, P.D. & Zimmerman, W.

Contributor/s

Justification
Previously listed as Extinct in the Wild from the 1960s up to the assessment in 1996. The species was then reassessed as Critically Endangered due to at least one surviving mature individual in the wild. Successful reintroductions have qualified this species for reassessment. The population is currently estimated to consist of more than 50 mature individuals free-living in the wild for the past five years. This taxon is threatened by hybridization with domestic horses, loss of genetic diversity, and disease. As the population size is small, it is vulnerable to stochastic events such as severe weather, Equus ferus przewalskii qualifies as Endangered under criterion D.

History
  • 2008
    Critically Endangered
    (IUCN 2008)
  • 2008
    Critically Endangered
  • 1996
    Extinct in the Wild
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Population

Population

There are now approximately 306 free-ranging reintroduced and native-born Przewalski’s Horses in Mongolia (Zimmerman 2011). All Przewalski’s Horses alive today are descended from only 13 or 14 individuals, which were the nucleus of a captive breeding program (Bowling and Ryder 1987). Introgression of domestic horse blood happened not only in Halle (#229 dom.Mongol), but also in Askania Nova (#175 Domina; Bowling et al. 2003).

Between 1992 and 2004, 90 captive-born horses were transported to the Takhin Tal/Gobi B reintroduction site in Mongolia (ITG International Takhi Group, Zimmermann 2008). A further three males were translocated from Hustai National Park to Takhin Tal in 2007 (Zimmermann 2008). In 2008 there were approximately 111 free-ranging horses in this population (Zimmerman 2008, Kaczensky and Walzer 2007). In December of 2009 there were 137 individuals in the population, but due to an extremely harsh winter (dzud) the population suffered extreme mortality and by August 2010 only 49 individuals remained (Kaczensky et al. 2010, Zimmerman 2011). From 1992 to 2000, 84 horses were brought to Hustai National Park by the Foundation for the Preservation and Protection of the Przewalski Horse and Mongolian Association for Conservation of Nature and the Environment (MACNE) from reserves in Europe (King and Gurnell 2005). As of the end of 2010 this population was approximately 233 individuals (Zimmerman 2011). A third reintroduction site was started in 2004 at Seriin Nuruu in the Khomiin Tal buffer zone of the Khar Us Nuur National Park in western Mongolia (Association pour le cheval de Przewalski: TAKH). Twenty-two individuals consisting of four pre-established families and one male bachelor group were brought from Le Villaret, France between 2004 and 2005 (C. Feh pers. comm., Zimmermann 2008). In 2010, this population had 24 individuals (Zimmermann 2011).

For the reintroduced population in Mongolia, mature individuals are those that are born in the wild and five years of age. However, individuals born in captivity do not count as mature until they have reproduced in the wild and that offspring is at least five years old. As of 2006 there were 55 mature individuals in the wild (52 (M.F., 26.26) in Hustai, 3 (1.2) in Takhin Tal). In 2007 Hustai had 68 (33.35) mature individuals and Takhin Tal had 11 (3.8) for a total of 79. The reintroduced populations continued to grow and in 2008, Hustai had 90 (39.51) and Takin Tal had 14 (7.7) mature individuals for a total of 104. In 2009, Hustai had a population of 118 mature individuals (52.66) and Takin Tal had 33 (15.18) for a total of 151. The winter of 2009/2010 was very severe and there was high mortality of Przewalski’s Horses, particularly in Takin Tal. In 2010, Hustai’s mature population was 117 (53.64) and Takin Tal’s number of mature individuals was reduced to 17 (8.9). However, the total population of mature individuals was 134. Hence for a period of five years, the mature population of Przewalski’s Horses in Mongolia has been more than 50 individuals. Although this means that the Przewalski’s Horse qualifies as Endangered it should be borne in mind that most of these individuals are from one reintroduction site and climatic perturbations like the extremely harsh winter in 2009/2010 can have very negative effects on small populations.

In China, the Wild Horse Breeding Centre (WHBC) of the Department of Forestry at Kalameili Nature Reserve (KNR) in Xinjiang Uighur Autonomous Region has established a large captive population of approximately 123 Przewalski’s Horses (January 2008, Pantel et al. 2006, Zimmermann et al. 2008). Since 2007 one harem group is roaming free on the Chinese side of the Dzungarian Gobi (Xinjiang); another 60 horses are roaming free during summer time but are returned to the acclimatization pen during the winter (Zimmermann et al. 2008).

The history of population estimates and trends in Przewalski’s Horse has been described by Wakefield et al. (2002). Since the ‘rediscovery’ of the Przewalski’s Horse for western science, western zoos and wild animal parks became interested in this species for their collections. Several long expeditions were mounted to catch animals. Some expeditions came back empty-handed and some had only seen a glimpse of wild Przewalski’s Horses. It proved difficult to catch adult horses, because they were too shy and fast. Capture of foals, with possible killing of the adult harem members, was considered the only option (Bouman and Bouman 1994). Four expeditions that managed to catch live foals took place between 1897 and 1902. Fifty-three of these foals reached the west alive. Between the 1930s and the 1940s only a few Przewalski’s Horses were caught and most died. At least one mare was crossbred with domestic horses by the Mongolian War Ministry (Bouman and Bouman 1994).

Small groups of horses were reported through the 1940s and 1950s in an area between the Baitag-Bogdo ridge and the ridge of the Takhin-Shaar Nuruu (which, translated from Mongolian, means ‘the Yellow Mountain of the Wild Horse’), but numbers appeared to decline dramatically after World War II. The last confirmed sighting in the wild was made in 1969 by the Mongolian scientist N. Dovchin. He saw a stallion near a spring called Gun Tamga, north of the Takhin-Shaar Nuruu, in the Dzungarian Gobi (Paklina and Pozdnyakova 1989). Annual investigations by the Joint Mongolian-Soviet Expedition have since failed to find conclusive evidence for their survival in the wild (Ryder 1990). Chinese biologists conducted a survey in northeastern Xinjiang from 1980 to 1982 (covering the area of 88–90°E and 41°31'–47°10'N) without finding any horses (Gao and Gu 1989). The last native wild populations had disappeared.

The number of living animals in the International Studbook was 1,872 in early 2008. Of the 53 animals recorded in the Studbook as having been brought into zoological collections in the west, only 12 contributed any genes to the current living population. Of these, 11 were brought into captivity between 1899 and 1902 and the last of them died in 1939. The twelfth founder was captured as a foal in 1947. The thirteenth founder was born in 1906 in Halle (Germany) to a wild-caught stallion and a domestic Mongolian mare, and the fourteenth founder is a female born in Askania Nova (Ukraine) to a Przewalski’s Horse stallion and a domestic female of a Tarpan type. Nevertheless, the current population is genetically very close to the original wild horses (Bowling et al. 2003). In addition to animals held in captivity and those already re-introduced, there have been a number of animals released into very large enclosures (reserves). The four largest are in Le Villaret (18.13; Massif Central, France), Buchara (19.17.1; Uzbekistan), the Hortobágy-National Park (77.81; Hungary), and the Chernobyl exclusion zone (32.37; Ukraine) (information as of January 2010, W. Zimmermann pers. comm.).

Population Trend
Increasing
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Threats

Major Threats
A number of causes have been cited for the final extinction of Przewalski’s Horses in Mongolia and China. Among these are significant cultural and political changes (Bouman and Bouman 1994), hunting (Zhao and Liang 1992, Bouman and Bouman 1994), military activities (Ryder 1993), climatic change (Sokolov et al. 1992), and competition with livestock and increasing land use pressure (Sokolov et al. 1992, Ryder 1993, Bouman and Bouman 1994). Capture expeditions probably diminished the remaining Przewalski’s Horse populations by killing and dispersing the adults (Van Dierendonck and de Vries 1996). The harsh winters of 1945, 1948, and 1956 probably had an additional impact on the small population (Bouman and Bouman 1994). Increased pressure on, and rarity of waterholes in their last refuge should also be considered as a significant factor contributing to their extinction (Van Dierendonck and de Vries 1996).

For the reintroduced populations, hybridization with domestic horses is the primary threat, accompanied by competition for resources with domestic horses and possibly other livestock. Wherever Przewalski's Horses come into contact with domestic horses, there is a strong risk of hybridization and transmission of diseases. Recently, illegal mining in the protected areas is an additional threat to the viability of these areas. In Hustai National Park, it has been noted that overgrazing of the buffer-zone and continued pressure on the reserve are possible consequences of the enhanced economic activity in this area (Bouman 1998); however, the second phase of the project (1998-2003) paid much more attention to sustainable development of the buffer-zone. In the western section of the Gobi National Park (Gobi B), habitat degradation by nomads and military personnel and their livestock continues; there is no core zone here that is free from human influence all year round. Infectious diseases transmitted from domestic horses, notably Babesia equi, B. caballi and strangles (infection by Streptococcus equi), are a major threat to small reintroduced populations originating from zoos (Roberts et al. 2005, King and Gurnell 2005). Predation on foals by wolves may account for a significant number of mortalities and constitutes a threat to the population growth and continued survival of this taxon (Wit and Bouman 2006, Kaczensky et al. 2004, Kaczensky and Walzer 2007). As was observed during 2009/2010, severe winters can result in significant mortality.

There is concern over loss of genetic diversity after being reduced to a very small population and maintained in captivity for several generations. Sixty per cent of the unique genes of the studbook population have been lost (Ryder 1994). Loss of founder genes is irretrievable and further losses must be minimized through close genetic management. Furthermore, inbreeding depression could become a population-wide concern as the population inevitably becomes increasingly inbred (Ballou 1994). However, correct management of the population can slow these losses significantly, as has been achieved since the organization of the regional captive-breeding programs.
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Management

Conservation Actions

Conservation Actions

Przewalski's Horse is legally protected in Mongolia. It is protected as Very Rare under part 7.1 of the Law of the Mongolian Animal Kingdom (2000). Hunting has been prohibited since 1930, and the species is listed as Very Rare under the 1995 Mongolian Hunting Law (MNE 1996). It is listed as Critically Endangered in both the 1987 and 1997 Mongolian Red Books (Shagdarsuren et al. 1987, MNE 1997), and in the Regional Red List for Mongolia (Clark et al. 2006). The taxon's entire reintroduced range in Mongolia is within protected areas. It is listed on CITES Appendix I (as Equus przewalskii).

The following conservation measures are in place:

  • An International Studbook was produced in 1959, followed in the 1970s by establishment of the North American Breeders Group, which developed into the Species Survival Plan for the Przewalski’s horse. The European Endangered Species Program for this species was accepted in 1986. Many countries now cooperate in these programs to minimize inbreeding and retain genetic diversity in their horse populations.
  • There are three ongoing reintroduction sites in Mongolia.
  • The Status and Action Plan for the Przewalski's Horse (Equus ferus przewalskii) was produced in 2002 (see Wakefield et al. 2002), and provides a more detailed account of the history and ongoing conservation efforts surrounding the species.
  • All three reintroduction sites are fully monitoring their populations and are integrating community livelihood support into their projects.
  • There have been several workshops of stakeholders involved in the reintroduction of Przewalski's Horse to Mongolia (Boyd 2009). At the ‘Endangered Wild Equid Workshop’ held in Ulaanbataar in 2010 the following threats were identified: loss of population due to stochastic events (i.e., severe winter); limited habitat and resources (pasture and water); domestic horses (hybridization, disease, social stress); lack of information, appreciation / awareness, lack of knowledge; and exploitation of resources (i.e., mining). Specific actions needed for each threat category were identified and described.

Conservation measures required:

  • The health of wild and domestic horses should be monitored for disease (Roberts et al. 2005). Standardized techniques should be used to monitor health, fecundity, mortality, habitat utilization and social organization of all populations (Wakefield et al. 2002), and contact between Przewalski's Horses and domestic horses should be kept to a minimum.
  • A single population management approach should be developed.
  • Mongolia currently has the only wild population and an action plan is needed for the country.
  • The genealogy of all horses in Mongolia should be established based on individual micro-satellite data to monitor inbreeding levels, identify hybrids and plan for necessary movements of horses between reintroduction centres to maximize genetic diversity.
  • An authoritative government protocol for hybrids should be developed, to be established before hybridization occurs, and to be made available in each re-introduction centre and to local people (King and Gurnell 2005).
  • Further communication and cooperation between all re-introduction centres would be beneficial.
  • Further training and post-graduate education of staff and biologists involved with this conservation work.
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Pleistocene Re-wilding

 

This species is one of a number which have been included in various “Pleistocene rewilding” plans. Pleistocene rewilding is the proposed practice of restoring ecosystems to their state in the Pleistocene, roughly 10,000 years ago. This contrasts the standard conservation benchmark, particularly in North America, of restoring ecosystems to their pre-Columbian or pre-industrial state. In both Eurasia and North America, the Pleistocene was characterized by much greater diversity and numbers of large herbivores and predators, including proboscidians, equids, camelids, and felidae (Donlan et al 2006; Zimov 2005). The process of restoration would involve the reintroduction of extant species in their historic range, as well as the introduction of ‘proxy organisms’ to replace the ecological functionality of extinct organisms (Donlan et al 2006). 

There are three central theoretical goals to Pleistocene rewilding. In Siberia, a team led by Sergey Zimov is investigating the role of large herbivores as ecosystem engineers. It is thought that herbivory pressure could play a central role in maintaining a grass-dominated plant community, as opposed to either tree- or moss-dominated. Grasslands are known to be more stable carbon sinks than either mossy or forested tundra, due to the rapidity of their biogeochemical cycling (Zimov 2005). In principle, then, reintroducing Pleistocene fauna could have positive climate change mitigation effects. Proposals in North America have focused instead on the preservation of ecological dynamics. Proponents of Pleistocene rewilding argue that due to the strong ecological interactions of megafauna, it is likely that their extinction at the end of the Pleistocene would have caused cascading ecological disruptions lasting until the present time (Donlan et al 2006). Additionally, introduction programs could provide a new lease on life for extant, endangered megafauna species, such as cheetahs and Asian elephants (Rubenstein 2006). 

Pleistocene rewilding, while headline-grabbing, is by no means the standard of modern conservation biology. There are a number of objections to the proposals of Pleistocene rewilders, summarized by Rubenstein et al (2006). The introduction of species which have been locally extinct for thousands of years, and more particularly the introduction of modern relatives of extinct species, carries many risks: the potential for invasive species, catastrophic disruption of existing ecosystems, inadvertent introduction of disease organisms, and unpredictable behavior of introduced species. Additionally, while paleoecology is a growing field, there is still a fair amount of uncertainty about the actual ecosystem functions of the Pleistocene.

Species which Zimov and his colleagues in Siberia are experimenting with bison, musk oxen, Przewalski’s horse, and Siberian tigers (Zimov 2005). Small-scale introductions have already begun in Yakutia. Donlan et al propose introducing Przewalski’s horse, Bolson tortoises, Bactrian camels, cheetahs, lions, and elephants into the Western United States (Donlan et al 2005). While some individuals of these species are present on privately owned land, there are no free-living populations in North America at this time. 

  • Donlan CJ, Berger J, Bock CE, Bock JH, Burney DA, Estes JA, Foreman D, Martin PS, Roemer GW, Smith FA, Soule ME, Greene HW. 2005. Pleistocene Rewilding: An Optimistic Agenda for Twenty-First Century Conservation. The American Naturalist 168:660-681.
  • Donlan, CJ. 2005. Re-Wilding North America. Nature 436:913-914.
  • Rubenstein DR, Rubenstein DI, Sherman PW, Gavin TA. 2006. Pleistocene Park: Does Rewilding North America Represent Sound Conservation for the 21st Century? Biological Conservation 132:232-238.
  • Zimov, SA. 2005. Pleistocene Park: Return of the Mammoth’s Ecosystem. Science 308:796-798.
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