ISSN 1028-334X, Doklady Earth Sciences, 2017, Vol. 477, Part 1, pp. 1247–1250. © Pleiades Publishing, Ltd., 2017. Original Russian Text © N.P. Kalmykov, D.V. Kobylkin, 2017, published in Doklady Akademii Nauk, 2017, Vol. 477, No. 1, pp. 57–60.

GEOLOGY

The First Find of the Cave Hyena (Crocuta crocuta spelea) on the Oka Plateau (Eastern Sayan)

N. P. Kalmykova,* and D. V. Kobylkinb Presented by Academician G.G. Matishov June 4, 2015

Received June 4, 2015

Abstract—These data indicate that the last representatives of Hyaenidae on the Oka plateau, i.e., cave hyenas (C. c. spelaea), are direct descendants of the hyenas that lived in the Altai–Sayan and Baikal mountain coun- try from the Late Miocene to the end of the Pleistocene. Their continued existence over a long time undoubt- edly indicates the absence of repeated glaciation during the Pleistocene. Cave hyenas are relics of the late Ter- tiary fauna in the area of Lake Baikal; they became extinct due to gradual cooling, which peaked at the end of the Late Pleistocene.

DOI: 10.1134/S1028334X17110010

Eastern Sayan is an asymmetrical fold structure of the Dibi River). The mummies belong to Plecotus sp., the Altai–Sayan mountain country adjacent to the Myotis sp., Eptesicus nilssonii, Canis lupus, Vulpes vul- southwestern margin of the Siberian Platform. It pes, and Moschus moschiferus, which inhabit Eastern extends from the Yenisei River in the northwest to the Sayan at the present time. southern edge of Lake Baikal in the southeast. The Oka Plateau or Central Sayan Upland is situated in the In addition, in this cave, the fossil remains of a cave central lower part of the structure; the absolute eleva- hyena—Crocuta crocuta spelaea (Fig. 2)—were found among the rock fragments. This finding of an almost tions are 2500–2600 m above the sea level. The pla- complete skeleton and the absolute age of the fossil teau is composed of volcanic–sedimentary and sedi- remains radically change existing hypotheses concern- mentary formations of the Late Riphean, Vendian, ing the migration of spotted hyenas from Africa, the and Lower and Middle Cambrian, which are generally time of their first occurrence, subsequent distribution, represented by limestone and dolomite. The following and extinction in the southern part of eastern Siberia. karst relief forms, new to the plateau, were revealed in This is the first find of a cave hyena on the Oka Pla- recent years: pillars, grottos, ponors, sinkholes, pits, teau, where, in the Late Pleistocene, the area of hyena arches, and caves [1]. Until recently, no such land distribution reached the upper boundary of the middle forms were known in the area of the Altai–Sayan altitudes; in the Altai–Sayan folded area, the fossil mountain country (Fig. 1), which is characterized by remains refer to the lower levels [2]. In Eurasia, all wide variety of karsting rocks—from Lower Protero- Pleistocene hyenas, which are attributed to the Cro- zoic marble to Ordovician red calcareous conglomer- cuta genus [4], are considered as subspecies of the ates [2]. Fossils and partial fossils of mammals [3] and spotted hyena, С. c. spelaea (cave hyena). However, their mummies have been found in Gorome Cave in some researchers consider them as a specific species— limestone of the Nyurgata and Khyuten formations of C. spelaea—which would certainly increase the numbers the Bokson series of the Cambrian exposed in the of this species. A 14С date of 26180 ± 810 (LU-7588) or middle reaches of the Zabit River (a left tributary of 30940 ± 700 cal yr BP was obtained for the fossil remains of the cave hyena in the Laboratory of Geo- chronology and Paleogeography, St. Petersburg State aInstitute of Arid Zones, Southern Science Center, University. The absolute age of the hyena remains Russian Academy of Sciences, Rostov-on-Don, 344006 Russia allows us to state that a hyena lived in Eastern Sayan at bSochava Institute of Geography, Siberian Branch, the final stage of the Karginsky Interglacial time Russian Academy of Sciences, Irkutsk, 664033 Russia (marine isotope stage MIS 3), the range of which is *e-mail: [email protected] 59000–24000cal yr BP.

1247 1248 KALMYKOV, KOBYLKIN

Uda R. K ° i 56 re n g N a

R

.

KanK R. a n R . Lena R.

Iya R.

Oka R. RRUSSIAU S S I A

Lake Baikal AreaA r e a studiedsoft u Studyd i e d Irkutsk 52° B.B. YYeniseienisei RR.. Irkut R. Ulan-UdeUlan-Ude

Selenga R. KhubsugulKhubsugul LLakeake

Orkhon R. MONGOLIAM O N G O L I A Ider R.

UUlanlan BatorBator

96° E 102° 108°

Fig. 1. Location of Gorome cave.

Cave hyenas appear to be a late stage in the evolu- autochthonous forms that inhabited Altai–Sayan and tion of spotted hyenas on the periphery of the vast dis- Baikal mountain country before and after the assumed tribution area in Eurasia and Africa. Different views waves of migration from Africa and South Asia. In on the origin and prochoresis of hyenas are currently Eastern Sayan, the most ancient representatives of this expressed. According to one version [5], Pachycrocuta family were found in the Upper Miocene deposits of brevirostris appeared in Asia, from where it migrated to Tyva, in Taralyk Cher, where fossil remains of the Africa ~3.5 million years ago and to Europe ~1.6 mil- hyena Adcrocuta [8] were identified. The phylogenetic lion years ago; it inhabited Eurasia and Africa ~0.5 position of this genus is still being discussed. However, and 3.0 million years ago, respectively [6]. According it is possible that this species is an ordinary Crocuta, to another version [7], cave hyenas of Northern Eur- because it belongs to the Crocuta crocuta sister group asia are of an African origin and are phylogenetically [9]. At the end of the Miocene and the beginning of associated with the spotted hyenas (C. crocuta) of the Pliocene, crocutas also inhabited an adjacent area, Africa. The recolonization of Eurasia by the spotted Western Mongolia: (Altan-Teli) and hyenas of African origin occurred in three waves: ~3.0, Crocuta gigantea 1.0, and 0.3 million years ago [4]. Crocuta sp. (Oshi) [10]. The occurrence of a hyena in the Late Miocene is not contrary to the current view There are no available or new data on the fossil on the Asian origin of the Crocuta genus; however, Hyaenidae in the southern part of eastern Siberia to new finds of Hyaenidae could correct this point of confirm these assumptions, because Hyaenidae are view.

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Crocuta cf. licenti inhabited the southern part of Eastern Siberia in the Late Pleistocene; it was identi- fied from the red formations of Western Transbaikalia (Tologoi, Beregovaya). С. brevirostris (=Hyaena brevi- rostris sinensis) inhabited the Southeastern Baikal Region (Zasukhino) in the Early Pleistocene (Eo-plei- stocene). A cave hyena (Crocuta spelaea) inhabited Western Transbaikalia (the upper horizons of the mid- dle formation in Tologoi) in the Middle Pleistocene. 1 Later, its scattered remains were found in Upper Pleis- 5 cm tocene deposits and in the Late Paleolithic sites of primitive man in Eastern Sayan, the Baikal Region, and Western Transbaikalia [11]. According to the published data, cave hyenas inhabited the Eurasia 50000–37000 years ago; in Altai and eastern Asia, they were spread in the same age range—48 650–12380 ± 1840 and 42300–1940 ± 840 years ago, respectively [4]. They inhabited the Tunka Valley (Eastern Sayan) in age range of 41486– 40033 cal yr ВР [14] and the Oka Plateau, according 2 to our data, 30940 ± 700 years ago. These dates indi- cate that they became extinct earlier in the Altai– Sayan mountain country compared to Western Fig. 2. The skull (1) and the lower jaw (2) of the hyena Europe and the Far East; in China, they probably sur- (Crocuta crocuta spelaea) from the Oka Plateau (Gorome Cave, Late Pleistocene). vived to the Holocene. One of the reasons was the rather intercontinental inland position of this moun- tain area, where the climatic conditions were not so extreme. In the Early Pleistocene (>3.5 million years ago), The represented data show that the last representa- hyenas (Pachycrocuta ex. gr. pyrenaica) lived in West- tives of Hyaenidae in the Oka Plateau, cave hyenas ern Transbaikalia (Udunga). This genus, like Plioro- (С. c. spelaea), are direct descendants of the hyenas cuta, Percrocuta, and Chasmaporthetes [11], is a syn- that inhabited the Altai–Sayan and Baikal mountain onym for the genus Crocuta, which was widespread in country from the Late Miocene to the end of the Pleis- Eurasia and Africa in the Late Miocene and the Pleis- tocene. Their continued existence over a long time tocene; it currently inhabits Africa, south of the undoubtedly indicates the absence of repeated glacia- Sahara. The genera mentioned, like Adcrocuta and tion during the Pleistocene. Cave hyenas are relics of Plesiocrocuta, also cannot be considered valid due to late Tertiary fauna in the area of Lake Baikal; they the lack of clear differences from the genus Crocuta. became extinct due to gradual cooling, which peaked They represent specific individuals of polymorphic at the end of the Late Pleistocene. populations of crocutas from oryctocoenoses of differ- ent ages, removed from one another. Their description as new taxons of a different level resulted in increased ACKNOWLEDGMENTS inflation, thereby making the already systematics This work was supported by the Russian Founda- more complex. This is confirmed by the revision of the tion for Basic Research, project no. 13-05-00521. Pleistocene hyenas of China [12]: Hyaena licenti was classified as the genus Pachycrocuta; H. sinensis, H. zdanksyi, and С. с. sinensis became synonyms of REFERENCES Pachycrocuta sinensis; H. ultima, H. honanensis, and 1. A. B. Imetkhenov, D. V. Kobylkin, and O. N. Morozov, H. spelaea were classified as the genus Crocuta. Later, in Proc. Irkutsk Geomorphological Seminar. Readings in P. licenti and P. sinensis were transferred to the P. bre- Memoriam of N. A. Florensov (Inst. Earth’s Crust Sibe- rian Branch Russ. Acad. Sci., Irkutsk, 2010), pp. 165– virostris synonymity, and all species Crocuta, to the 166 [in Russian]. Crocuta crocuta synonymity [13]. Simplification of 2. R. A. Tsykin, in Caves (Perm, 1993), pp. 59–70 [in Rus- their taxonomic diversity, which has not still been sian]. drawn to its logical conclusion, assumes that in East- 3. D. V. Kobylkin, in Proc. All-Russ. Sci. Conf. with Inter- ern Asia the genus Crocuta existed from the Late Mio- national Participation “Problems on Wild Life Studying cene to the end of the Pleistocene, and the above- and Protecting in the North” (Komi Sci. Center Ural mentioned species likely appeared to be the successive Branch Russ. Acad. Sci., Syktyvkar, 2009), pp. 62–64 links of a certain phylogenetic line of crocutas. [in Russian].

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