Preliminary analyses of the frozen mummies of mammoth ( Mammuthus primigenius ), bison ( Bison...

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© 2013 International Society of Zoological Sciences, Institute of Zoology/ Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd

Integrative Zoology 2014; 9: 471–480 doi: 10.1111/1749-4877.12079

ORIGINAL ARTICLE

Preliminary analyses of the frozen mummies of mammoth (Mammuthus primigenius), bison (Bison priscus) and horse (Equus sp.) from the Yana-Indigirka Lowland, Yakutia, Russia

Gennady G. BOESKOROV,1 Olga R. POTAPOVA,2 Eugeny N. MASHCHENKO,3 Albert V. PROTOPOPOV,4 Tatyana V. KUZNETSOVA,5 Larry AGENBROAD2 and Alexey N. TIKHONOV6

1Diamond and Precious Metal Geology Institute, Siberian Branch of Russian Academy of Sciences, Yakutsk, Russia, 2The Mammoth Site of Hot Springs Museum, Hot Springs, South Dakota, USA, 3Paleontological Institute, Russian Academy of Sciences, Moscow, Russia, 4Department of the Mammoth Fauna Studies Academy of Sciences of the Sakha (Yakutia) Republic, Yakutsk, Russia, 5M. V. Lomonosov Moscow State University, Moscow, Russia and 6Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia

AbstractThe frozen bodies of a young woolly mammoth (Mammuthus primigenius), a wild horse (Equus sp.) and a steppe bison (Bison priscus) were recently found in the northern Yakutia (northeastern Siberia). All specimens have preserved bones, skin and soft tissues. Whereas the woolly mammoth and the Pleistocene horse were rep-resented by partial frozen bodies, the steppe bison body was recovered in an absolutely complete state. All spec-imens were found frozen in the permafrost, with some of the tissues mummified. The wild horse and steppe bi-son are of Holocene age, and the mammoth is of Late Pleistocene age.

Key words: bison, horse, mammoth, mummy, Siberia

Correspondence: Alexey N. Tikhonov, Zoological Institute, Russian Academy of Sciences, University embankment 1, St. Petersburg 199034, Russia. Email: [email protected]

INTRODUCTION The special climatic and geological conditions that

have prevailed in the Yakutia Republic (Eastern Siberia) for thousands of years have enabled the preservation of the unique ancient permafrost deposits. Formed during the extremely cold periods of the Pleistocene glacia-tions, the permafrost deposits, also referred to as the ‘ice

complex’, remain intact due to the cold climate that has been dominant in northeastern Eurasia.

The permafrost in Yakutia territory is famous for finds of ‘graveyards’. Skeletons, numerous isolat-ed bones and even frozen corpses of Ice Age mammals have been discovered. Most of those finds were locat-ed in the arctic region of Yakutia, where icy depos-its of the Late Pleistocene Edoma Formation are wide-ly distributed. Since the first sensational paleontological finds of the complete carcasses of the Adams (1799) and Berezovka (1901) mammoths, many frozen carcass-es and partial carcasses of Ice Age mammal specimens have been found in the northern Yakutia: Berelekh wol-verine (1970); Kolyma woolly rhinoceros (2007); Sel-

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erikan (1968) and Dyukarski horses (1981); Mylakh-chin (1971), Yana (2009) and Tsiigehtchic steppe bison (2007); Sanga-Yuryakh (1908), Lyakhov (1908, 1993, 1995), Berelekh (1970), Shandrin (1972), Maksunuo-kha (1997, 2000) and Yukagir (2002) mammoths; Abyi (1990), Oymyakon (2002) and Khroma (2009) mam-moth calves; and Yukagir mammoths (Vereshchagin & Lazarev 1977; Vereshchagin 1979; Vereshchagin & Tik-honov 1999; Lazarev et al 2001; Lazarev 2002; Boe-skorov et al. 2007; Zazula et al. 2009; Boeskorov et al. 2011).

During the study of the preserved carcasses, a large amount of scientific information have been obtained, including on anatomy, morphology, animals’ precise age, nutrition state at time of death, causes of death and paleoecological conditions of the areas (Zalens-ki 1903; Flerov 1931; Heintz 1958; Ukraintseva 1977, 1993; Vereschagin & Lazarev 1977; Ukraintseva et al. 1978; Lazarev 1980, 2008; Ivanova 1981; Vereschagin 1981; Dubrovo 1982; Guthrie 1982; Shilo et al. 1983; Maschenko et al. 2005; Mol et al. 2006; Boeskorov et al. 2007, 2010, 2011; van der Geel et al. 2008, 2011; Boeskorov et al. 2011; Kosintsev et al. 2011; Maschen-ko et al. 2012a). This recently discovered material has allowed application of new methods that have never been used in studies of mummies of extinct species be-fore (Fisher & Beld 2002; Rountrey et al. 2010, 2011; Shirley et al. 2011). Frozen cadavers, along with other mammal remnants from the Siberian permafrost, have attracted the attention of microbiologists and molecular genetics. As a result, the woolly mammoth mitochon-drial DNA has been successfully sequenced (Debruyne et al. 2003; Poinar et al. 2006; Rogaev et al. 2006; Gil-bert et al. 2007, 2008) and completion of the nuclear ge-nome decoding of this species is making good progress (Miller et al. 2008).

MATERIALS AND METHODSThe primary research of the specimens was carried

out in Mar 2012 by the international team from Rus-sia (Yakutsk, Moscow and St. Petersburg), USA, France and Japan.

The frozen corpses were transported from the field storages to Yakutsk in 2011–2012, for thorough study in stable conditions at Diamond and Precious Metal Geol-ogy Institute. All specimens were partially thawed out to allow morphological measurements, hair, tissue, stom-ach and intestine content sampling, detailed morpholog-ical descriptions, photography and documentation.

The Yuka mammoth corpse included approximate-ly 95% of its hide, and soft tissues around limbs were preserved in articulated position. The skull, mandible, axis, both scapula, 8 ribs, pelvis with sacrum and tail bones were found near the body. The Yuka mammoth condition did not allow the use of standard methods for measuring modern elephants’ shoulder heights (Whyte 1996) because large portions of the skeletal elements were extracted from the body before the study. Instead, we used the approach for measuring mammoth carcass-es suggested by Vereschagin (1981).

The Yukagir bison corpse included 100% of the body. At the time of the present study its skeleton was not yet available for study and measurement. The bison was ex-amined following the method for measuring the Euro-pean bison skin outlined by Sokolov (1979). The body measurements were taken with large calipers and a tape. Due to the ‘locked’ position of the bison body (see de-scriptions below), we could not use the standard mea-surements for Bovidae and domestic cattle but de-veloped original measurements that fit this particular specimen (overall body, head, horn cores, and front and hind legs). Preliminary determination of the age of the bison specimen was based on the incisors eruption and wear (Fuller 1959; Frison & Reher 1970).

The Yukagir horse was represented by 2 parts of the body, the head with the neck, and a hind part with both legs. The age of the specimens was determined based on the exposed lower incisors eruption and wear (Liskun 1949) and general measurements were taken following the approach of Gromov et al. (1963).

History, geography and geology of the finds

Between 2009 and 2011 the following unique speci-mens were discovered from Yakutia: the frozen body of a young mammoth [Mammuthus primigenius (Blumen-bach, 1799)], a wild horse (Equus sp.) and a steppe bi-son (Bison priscus Bojanus, 1827).

In the summer of 2009, on the southern coast of the Dmitry Laptev Strait in the area called Oyagossky Yar, 30 km west from the mouth of the Kondratieva River, the frozen remains of a young mammoth (Yuka) were found. In Aug the following year, just 50 m from the ‘Yuka site’, a partial frozen body of a horse was discov-ered. In mid-Aug 2011, a complete frozen corpse of a steppe bison was found on the northwest shore of the Lake Chukchalakh in the northern part of the Yana-In-digirka Lowland. The place of its discovery is approxi-mately 100 km southwest of the ‘Yuka mammoth site’.

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Frozen mummies from Yakutia, Russia


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All 3 finds were made by members of the tribal com-munity ‘Yukagir’ during their traditional summer activi-ties (hunting, fishing and commercial gathering of mam-moth tusks). Members of this community had found the remains of mammoths in the past. In 1994, on the bank of the Maksunuokha River they found a ‘graveyard’ of mammoths containing 160 bones belonging to sev-eral individuals, as well as 3 complete feet with mus-cles and skin from the same specimen (Boeskorov et al. 2006; Lazarev 2008). In 2002, sons of the communi-ty head Vasilii Gorokhov discovered parts of a Yukagir mammoth: the head with tusks, the front left leg, skin fragments, strands of wool and a few postcranial bones. Multidisciplinary studies of this mammoth are published as a separate monograph (Boeskorov et al. 2007). The Yukagir mammoth head is known, having been exhibit-ed at the 2005 World Expo in Nagoya, Japan and being the main logo for the expo.

The place of discovery of the bison’s frozen body is approximately 25 km north from the location of the Yu-kagir mammoth. There are no obvious differences be-tween these places in regards to the landscape (the east-ern shore of the Chukchalah Lake has an unnamed hill nearby, with elevation 558 m asl; the Yukagir bison site is located in its foothills). The deposits that make up this rolling hill landscape are dissected by the Maksunuokha River and covered by numerous lakes. The deposits are represented by the Late Pleistocene Edoma Formation. Radiocarbon dating of the Yukagir mammoth revealed that it lived approximately 18 400 years BP, during the maximum of the Sartan glaciation (Boeskorov et al. 2007).

The sites of the new finds are located in the Yana-In-digirka lowland, particularly between the Omoloy Riv-er valley on the west and the Indigirka River on the east, which represent some of the most remote, rugged and undeveloped regions of the far north of Yakutia. The lowland is heavily waterlogged, with numerous lakes, and composed of the icy Pleistocene silts of the Edoma suite (Kaplina 2011), covered with powerful polygonal ice wedges. In the northern Siberian lowlands the arctic tundra is widespread (Boeskorov et al. 2007). Ancient soils were recovered from the discovery sites in the tun-dra permafrost. A qualitative study of the pollen from the ancient buried soils of the Upper Pleistocene of the ‘Yukagir mammoth site’ attributed the overall vegeta-tion composition to steppe formation.

The Yana-Indigirka lowland, as a part of North-ern Eurasia in the Late Pleistocene, was represented by vast tundra-steppe biome composed of a large vari-

ety of grasses. This nutritious steppe component deter-mined a high number and variety of the Late Pleisto-cene mammalian fauna in the region, which along with the extremely favorable preservation conditions became a key factor for preserving complete specimens of ex-tinct animals. The Oyagossky Yar, where the carcasses of mammoth and horse were found, is one of the rich-est areas yielding Quaternary fossils in the northern Ya-kutia. It is located on the mainland coast of the Dmi-tri Laptev Strait and stretches more than 100 km from Cape Saint Nose (Svyatoi Nos) in the west to the mouth of the Kondratieva River in the east. The Edoma depos-its of the Oyagossky Yar’s northern slope can reach 40–50 m asl The formation of the icy deposits of this loca-tion includes the entire second half of the Pleistocene. The western part the Oyagossky Yar is composed of the lake-alluvial silt sediments reaching 5–7 m asl, which are contemporary with the sediments of the Olersky For-mation (Lower Kolyma River basin, Lower Pleistocene) (Tomirdiaro et al. 1982). The stratotype of the Oyagoss-ky Formation was described at this location. The low-er deposits, which were accumulated during the Zyryan glaciation of the Late Pleistocene in Siberia, produced a radiocarbon date of more than 47 000 years BP. The lay-ers of the upper unit of the type formation are dated be-tween 37 000 and 33 000 years BP (Karginsky intersta-dial) (Kaplina & Lozhkin 1982; Tomirdiaro et al. 1982). The eastern part of the Oyagossky Yar (the Kondratyeva River area) yielded bones of the Pleistocene mammals that were dated between 47 000 and 26 000 years BP (Kargin-sky interstadial) and between 17 000 and 16 000 years BP. (Sartansky stadial) (Nikolskyi & Basilyan 2003).

The studies of the Oyagosski Yar Pleistocene remains revealed the following fauna: Canis lupus Linnaeus, 1758, Alopex lagopus (Linnaeus, 1758), Gulo gulo (Lin-naeus, 1758), Ursus sp., Panthera leo spelaea (Gold-fuss, 1810), M. primigenius, Coelodonta antiquitatis (Blumenbach, 1807), Equus lenensis (Kahlke, 1994), Rangifer tarandus (Linnaeus, 1758), Cervus cf. elaphus (Linnaeus, 1758), Alces sp., B. priscus, Ovibos pallan-tis Blainville 1816 and Saiga borealis (Tschersky, 1876) (Tomirdiaro et al. 1982; Sablin 1991).

RESULTS AND DISCISSION

The Yuka mammoth

By tradition, most of the significant mammoth re-mains are given proper or geographical names to com-memorate the find and for easy recognition of the spec-imen: for example, Effie (Alaska, 1947), Dima (Dima

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Creek, Magadan District, 1977), Masha (Yamal Penin-sula, 1988) and Luba (Yamal Peninsula, 2009). In ac-cordance with this tradition, the young mammoth found was named ‘Yuka’, in honor of the community ‘Yukagir’, named after the ‘Yukagir’ tribe currently inhabiting the area.

The Yuka mammoth (field number: OYu No. 1–13 is the property of the municipal community ‘Yukagir’, and is currently being studied and stored in monitored con-ditions in Yakutsk (Academy of Sciences of the Sakha [Yakutia] Republic).

Despite several injuries and cuts in the neck, back and shoulder areas, the skin (the texture and the surface layer) of Yuka is well preserved (Fig. 1). The hair is pre-served on the legs and both sides of the body. The tem-poral and mammary glands, left ear, trunk (with all 3 finger-like appendages), and upper and lower lips are in good condition. The trunk is intact, which is very rare for corpses of mammoths, due to predators and scav-engers. The majority of the limb bones were preserved in anatomical order. Soft tissues covered by skin were preserved from the distal ends of the humerus and fe-mur to the distal feet bones. Separate from the body, the skull, the lower jaw, first cervical vertebra, right and left shoulder blades, pelvis anatomically connected to the sacrum, last lumbar vertebra and caudal vertebras #1–3 and 9 ribs from the right side were found. All the sepa-rately preserved bones of the mammoth Yuka are cov-ered by a large amount of remnants of mummified soft tissues (muscles and ligaments). A part of the left hind foot (all bones and tissues below astragalus) is missing: it probably was eaten off by predators (most likely the polar fox). Traces of small canine punctures (possibly from the polar fox) are evident on the skin of the tail.

The special scientific value of Yuka is that this mam-moth appears to be a very young individual (approxi-mately 8–10 years old), based on the dental evidence (DP4/M1). Among known complete mammoth speci-

mens, the Yuka mammoth is unique due to its age. This find filled one of very few remaining ‘blank spots’ in our knowledge of individual development of mammoth at this age, with its growth rate and main characteristics showing mammoth adaptation to the cold climate of the Ice Age.

The morphology of the external genitalia remnants was sufficient to indicate that the mammoth Yuka is a female. The shoulder height of the mammoth did not ex-ceed 165 cm, the length from the front of the forehead (the base of the trunk) to the base of the tail measured approximately 205 cm, and its ‘live’ weight is estimated at 450–550 kg. It was already a semi-adult animal that could live independently as a part of a mammoth family group.

One of the unexpected characteristics of this spec-imen (of this age) was the presence of the permanent tusks ‘erupted’ from the alveoli, but their front ends had not yet reached the surface of the skin. Another unusual feature was the presence of 4 nails on each of the front feet (Fig. 2). In other woolly mammoth specimens, front feet usually possess 3 nails.

As a majority of the radiocarbon dates received from other mammal species have been collected from the Oyagossky Yar, Yuka lived (and died) in the Kargin-sky interstadial (C14–34 300 ± 250 years BP (39 440–38 850 calBP; GrA-53289) (Boeskorov et al. 2012). The cause of the mammoth Yuka death has not yet been determined; however, there are 2 types of skin damage on the body. The skin on the shoulder and throat, the left side of the back and the inner side of both hind legs contains deep scratches and cuts, corresponding to the

Figure 1 The Yuka mammoth.Figure 2 The soles with visible hoofs on the forefoot of the Yuka mammoth.

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claw sizes of a very large predator, possibly a cave lion, P. spelaea. However, the wounds did not penetrate the skin deep enough to have directly caused the death of the mammoth. Other significant skin damage consists of large missing portions of the hide, on the neck and on 1 side of the body. The hide cuts have a repeating ‘ladder’ pattern, which is not compatible with any known cut mark pattern inflicted by the Paleolithic artifacts discov-ered in the Eastern Siberia (Maschenko et al. 2012b).

The Yukagir horse

The partial carcass of the horse represents the re-mains of an adult mare approximately 5 years old. The frozen corpse is represented by a well-preserved head and neck (detached from the body) (Fig. 3), and the hind part of the body with internal organs in the abdom-inal cavity, hind legs and tail (Fig. 4). The front legs are missing. Apparently, this horse fell victim to predators;

the preserved areas of the skin contain many punctures, holes and scratches comparable to claw sizes of large predators. The upper part of the snout (frontal maxilla) is missing; it was probably was eaten off together with the front legs and the rest of the torso. The skin covers most of the head area, the neck, the back and the sides of the body, and completely covers the hind legs. Mum-mified skin is hardened, and for the most part has a dark brown color.

Body measurements show that the height of the horse at the hindquarters was approximately 130 cm. Thus, the new finding relates to the ‘undersized’ horse, simi-lar to the Lena horse (E. lenensis), which inhabited the Eastern Siberia (Gromov & Baranova 1981) and proba-bly went extinct during the Holocene. The radiocarbon date of the Yukagir horse is 4630 ± 35 years BP (GrA-540209, CalPal calibration: 5384 ± 58 calBP) (Boeskor-ov et al. 2012), which makes this find one of the latest representatives of the extinct species in the Palearctic. Further studies will determine the horse’s species affil-iation and establish whether it could be the evidence of the species survival into the Holocene, in the northern Yakutia.

Mummified remains of the Pleistocene horses, in-cluding Selerikan, Dyukarsk and Batagai horses of Ya-kutia (Vereshagin & Lazarev 1977; Lazarev 2002, 2008), Bilibino horses from Chukotka Peninsula (Spass-kaya et al. 2012), the isolated limbs of horses from the Easter and Goldstream creeks, Alaska (Guthrie 1990), and partial frozen and mummified Yukon horse Equus lambei Hay, 1917) remains from last Chance Creek, Yu-kon (Harington & Eggleston-Stott 1996), are stored and exhibited in museums in Russia, the USA and Canada. The scarcity of the mummified horse remains highlights the high scientific and museum value of the new discov-ery.

The Yukagir bison

This bison constitutes a unique find because the com-plete and intact frozen body of the adult animal has been preserved (Fig. 5). It is the third reported complete finding of the species since the discovery of the ‘Blue Babe’ from Alaska (Guthrie 1990) and the Mylakh-chin bison from Yakutia (Lazarev 1980), and the first complete specimen of the species from the Holocene age. Mitochondrial DNA analyses of the hair revealed that it belongs to a steppe bison, B. priscus (Boeskor-ov et al. 2012). It has all organs fully preserved, includ-ing the snout, the tips of the ears, the tail, the genitalia, and contents of the stomach and intestines. The bison is

Figure 3 The head of the Yukagir horse.

Figure 4 The hind part of the Yukagir horse carcass.

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a male of approximately 4 years. The distance between the horns tips is approximately 75 cm, the height at the withers of the animal was approximately 170 cm and the weight is estimated to have been approximately 500–600 kg. The spread of horn cores places this specimen within the range of the Late Pleistocene (Wurm–Holo-cene) B. priscus occidentalis from East Siberia (670–800 mm) (Flerov 1977), which was smaller than its larger predecessor in Siberia Bison priscus crassicoris Rich-ardson, 1854 (Flerov 1977). The latter form is some-what smaller than the North American Late Pleistocene contemporaneous species Bison occidentalis Lukas, 1898 (mean 779 cm; minimum/maximum 626–1055 cm) and significantly smaller than the Bison antiquus Leidy, 1852 (mean 870 cm; minimum/maximum 765–1067 cm), the descendant of B. latifrons (Harlan, 1825) (Skinner & Kiesen 1947; McDonald 1981), all from North America. However, the relationship between the East Beringian B. occidentalis and B. antiquus remains unresolved (Wilson et al. 2008). The spread of the horn cores of the Yukagir bison is also considerably smaller than that of the Tsiigehtchic steppe bison from Macken-zie River valley, Yukon, with radiocarbon age of 11 830 ± 45, years BP (940 mm) (Zazula et al. 2009), Blue Babe from Alaska with radiocarbon date approxi-mately 36 000 years BP (922 mm) (Guthrie 1990), and fossil male bison from interior Alaska (mean 986 mm) (Guthrie 1990).

The bison mummy was found in a ‘sleeping pose’, with legs tucked under the belly, the neck stretched out and the head lowered to the ground: a typical position for ungulates that have died naturally. Moreover, this natural death is evidenced by absence of any damage from predators.

Fossil remains of the complete bison mummies are rare. Before the Yukagir bison find, 2 other permafrost mummies were known. The famous ‘Blue Babe’ from Alaska that is exhibited as a stuffed, reconstructed ani-mal in the Museum of the University of Alaska in Fair-banks was the most complete until now (Guthrie 1990). Another frozen mummy of the steppe bison was discov-ered in the basin of the Indigirka River and named ‘My-lakhchin bison’ (Lazarev 2008). However, the preserva-tion of the Yukagir bison is incomparably better than the latter. Radiocarbon dating of the horn core and hair sam-ples yielded very close results (with Groningen calibra-tion): C14 – 9310 ± 45 years BP (10 575–10 435 calBP [1950]; GrA-53290, horn core), and C14 – 9295 ± 45 BP (10 570–10 425 calBP [1950], GrA-53292, hair) (Boeskorov et al. 2012).

The value and aspects of investigation of these unique finds

The cadavers of 3 specimens were sampled for mi-crobiological, radiocarbon, palynological, molecular ge-netic (DNA), lithological and other types of analysis, to allow, for example, identification of the time of the death of animals, their nutrition and the composition of vegetation in the period of their existence. Their genet-ic characteristics (relationship to closely-related species and forms), the taphonomy of the sites and the cause of the animals’ death will also be studied. These analyses to reconstruct the paleoecology of these extinct animals will be undertaken in the near future.

An important aspect of the research will be computed tomographic scanning and a related study of isotopes of nitrogen, carbon and oxygen from the teeth of animals, which is likely to be held in conjunction with an inter-national team from Russia, the USA and France. The study of changes in the composition of isotopes of these elements can allow identification of the timing of onto-genetic development, physiological features, diet and environment within a few days over the entire life of the animals, as well as the season of their death. Microbio-logical research of all objects will be performed to ex-plore paleomicro-flora and hidden high-risk infections. In Yakutsk, the lithological investigation of sediments from the burial places is currently being conducted to identify the site’s deposit types (the lacustrine-alluvi-al or others), which were formed under strictly defined conditions.

The sequencing of ancient DNA has become a tradi-tional subject of the studies applied to the extinct spe-cies. New data on the bison DNA may help resolve

Figure 5 The Yukagir bison.

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some uncertainties regarding the systematics of these animals, including the degree of relationship between the Pleistocene bison of Siberia with extinct and modern bison in North America. These data will be used to in-vestigate their relationship with the newly-described ex-tinction of populations of bison and their migrations in the western and eastern Beringia during the Late Pleis-tocene (Shapiro et al. 2004).

The ‘Yukagir’ horse may provide new information on the taxonomy and origin of the horses in far northeast-ern Siberia, including their relation to the modern Yaku-tian breed of horse. The Holocene dates for the horses from northern Siberia clearly show that the Pleistocene horse survived in Eastern Siberia until approximate-ly 2000–4000 years BP (Lazarev 1980; Kuznetsova et al. 2001; Boeskorov 2004), and the new find represents additional proof of this interesting phenomenon. Some Russian scientists (Cherskii 1891; Gabyshev 1957; Laz-arev 1980) have suggested that the Yakut domestic hors-es represent descendants of the Late Pleistocene species. However, data received from protein electrophoresis overruled this theory and confirmed the observation ex-pressed by Kuzmina (1977) and Boeskorov (2000) that the morphological similarity of the Late Pleistocene horse and the domestic Yakut horse could be the result of parallel (independent) adaptation to the cold climate. The present study also revealed that genetic continui-ty of the recent Yakutian horse is close to some of the small-sized Central Asian breeds of Equus caballus L. (Guriev 1998; Tikhonov et al. 1998).

All described finds are stored in frozen condition and will be dissected and sampled for different research projects in the future. Upon completion of research, the specimens will be exhibited in museums in the Yakutian Republic.

ACKNOWLEDGMENTSThe authors are grateful to the late Vasiliy Gorokhov,

the head of the tribal community ‘Yukagir’ (who passed away in Nov 2012), and to the Yukagir community for allowing us to borrow the specimens for this study in Yakutsk, Russia. The Scientific Research Institute of Applied Ecology of the North of Northeastern Feder-al University, Yakutsk and Bernard Buigues (Saint-Jean de Lus, France) helped with the logistics and provided funds for the specimens to be transported to Yakutsk in 2012.

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