Cadernos Lab. Xeolóxico de Laxe ISSN: 0213-4497 Coruña. 2001. Vol. 26, pp. 373-398 Pleistocene small cave bear (Ursus rossicus) from the South Siberia, Russia Un pequeño Oso de las Cavernas (Ursus rossicus) del Sur de Siberia, Rusia BARYSHNIKOV, G.1, FORONOVA, I.2 AB S T R A C T The skull, mandibles and cheek teeth of U. rossicus from four localities of the South Siberia are examined. This species inhabited the steppe regions in early Middle and Late Pleistocene. By odontological characters it is more close to U. r. rossicus from Krasnodar, than to U. rossicus uralensis from Kizel Cave in Ural. Discriminant analysis, based on measurements of lower cheek teeth of the cave bears from seven sites of Europe and Siberia, demonstrated that U. rossicus most resembles morphometrically U. savini. As a result of cladistic analysis employed 17 characters of skull, limb bones, and dentition, the phylogenetic tree has been obtained for 7 species of the genus Ur s u s . A four species of the cave bears are included in the subgenus Spelearctos: U. savini, U. rossicus, U. denin - geri and U. spelaeus. Key words: cave bears, Ur s u s , Siberia, Pleistocene, evolution (1) Zoological Institute, Russian Academy of Sciences, Universitetskaya naberezhnaya 1, 199034 St. Petersburg, Russia; e-mail: [email protected] (2) Institute of Geology, Siberian Branch of Russian Academy of Sciences, pr. Akad. Koptiuga 3, 630090 Novosibirsk, Russia; e-mail: [email protected] 374 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001) INTRODUCTION V E R E S H C H A G I N & T I K H O N O V, 1994; BA RY S H N I K O V , 1995). These Small cave bear Ursus rossicus Borissiak, findings are also associated with the zones 1930 has been widely distributed in of spreading of the Pleistocene grass com- Northern Eurasia in the Middle and Late munities. Pleistocene. In East Europe this species ALEXEEVA (1980), who gave brief was found in the south of Ukraine, characteristics of U. rossicus from Krasnyi Northern Caucasus, Lower Volga River, Ya r, came to the conclusion that the Bolshoi Irgiz River basin, Ural River and Siberian bear had been more specialized to in foothills of Ural Mountains consume plants than other geographic (BORISSIAK, 1932, VERESHCHAGIN races. In the later paper she (ALEXEEVA, 1959, 1982, K U Z M I N A, 1975, 1996) proposed a new subspecies name U. BARYSHNIKOV et al., 1991). It is also rossicus obensis, but did not give a diagno- known from the plain areas of Kazakhstan sis and did not designate the type, which (KOJAMKULOVA, 1969). Two subspe- makes this name not acceptible according cies were distinguished: U. r. rossicus from to the Code of Zoological nomenclature. Northern Caucasus (Krasnodar), Lower Vereshchagin & Tikhonov (1994) gave the Volga (Kopanovka) and adjacent plains, map of findings of the small cave bear in and U. r. uralensis Vereshchagin, 1973 Siberia. FORONOVA (1999, in press) from Kizel Cave and other cave localities published the photographs and brief mor- of Middle and South Ural. The geography phological description of mandibles of U. of fossil findings points out that the small rossicus from Kuzbass localities. The com- cave bear was mainly a dweller of open plete morphological analysis of the herbaceous areas including the steppe Siberian material is still absent. ones. It did not evidently penetrate to This paper presents the first detailed West Europe where its remains are not description of skull fragments and teeth of recorded. the small cave bear from the collection of In Siberia, N. Vereshchagin first iden- Zoological Institute Russian Academy of tified U . r o s s i c u s by os penis from Sciences in Saint Petersburg (ZIN) and Strashnaya Cave in Altai Institute of Geology (IG), and Institute of (OKLADNIKOV et al., 1973). It was elu- Archaeology and Ethnography (IAE), cidated later that small cave bear was dis- Siberian Branch of Russian Academy of tributed over the whole of South Siberia. Sciences in Novosibirsk. It was recorded at Irtysh River (Omsk Region), Ob River (Krasnyi Yar), Altai Mountains (Strashnaya Cave, Denisova MATERIAL AND METHODS Cave), Kuznetsk Basin (Bachatsk and The material studied contains the skull Mochovsky quarries), Yenisei River fragment (ZIN 32748) and the right man- (Kurtak) and in Transbaikalia (Tologoi on dible (ZIN 35075) from Krasnyi Yar in Selenga River) (A L E X E E VA, 1980; Tomsk Province, the right mandible (IG F O R O N O VA , 1982, 1999; 328) from Bachatsk quarry (figure 1) and CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 375 Figure 1. Right mandible of Ursus rossicus from Bachatsk quarry, IG 328. Lateral view. the left mandible (IG 10128) from and University of Helsinki), and from the Mokhovo quarry (figure 2) in Kemerovo Secrets Cave, Ural Mountain (our data). Province, 19 mandible fragments (IAE For the cranial characters analysis, we 36, 63, 81, 88, 89, 93, 122, 171, 235, examined also skulls of recent U. arctos L. 330, 331, 455 et al.) from Berezhekovo, and U. maritimus Phipps from ZIN collec- Kurtak archeological region, Krasnoyarsk tion. We used the information on U. etrus - Territory (figure 3, 4), and also several iso- cus G.Cuvier published by MAZZA & lated check teeth and limb bones from RUSTIONI (1992). Additional data on these localities. We have also used the cave bear was obtained from Andrews photograph of the skull from Kurtak, (1922), BORISSIAK (1932), RABEDER kindly provided by Dr. Nikolai Ovodov (1983), RABEDER & TSOUKALA (Novosibirsk). (1990), M A Z Z A et al. (1995), The material examined was compared Baryshnikov (1998), VERESHCHAGIN with the collections of Ursus savini & BARYSHNIKOV (in press). Andrews from Bacton Forest Bed and U. In the mode of measurements for skull deningeri von Reichenau from Westbury and mandible we follow von den Quarry Cave, England (Natural History DRIESCH (1976). Cheek teeth were mea- Museum, London) and Kudaro 1 Cave, sured following B A RY S H N I K O V Caucasus (ZIN), U. rossicus uralensis from (1998). Dimensions were taken with dial Kizel Cave, Ural (ZIN), U. spelaeus calipers with accuracy up to 0.1 mm. The Rosemüller from Arcy-sur-Cure, France data were processed by Factor Analysis, (Laboratoire díEthnologie prehistorique, Cluster Analysis and Discriminant Nanterre), from Odessa, Ukraine (ZIN Analysis from STATISTICA, 6.0. In 376 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001) Figure 2. Left mandible of Ursus rossicus from Mokhovo quarry, IG 10128. Lateral view. Figure 3. Fragment of right mandible Ursus rossicus from Berezhekovo 4, IAE 171. Occlusal view. Discriminant Analysis, we used the for- Pleistocene mammal fauna in the south of ward stepwise method. PAUP computer Western and Middle Siberia: Kuznetsk program (version 3.1.1) was applied for Basin, Krasnyi Yar at the Ob River, and the phylogenetic analysis. Kurtak archaeological region. The Kuznetsk Basin is situated in the GEOLOGICAL SETTING AND southeast of the Western Siberia. It was STRATIGRAPHY formed in the Late Cenozoic as an enor- mous intermountain depression, restricted The studied material is originated by the Kuznetsk Alatau, Gornaya Shoriya, from the famous localities of the and Salair Ridge. This region is one of the CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 377 Figure 4. Left mandible of juvenal Ursus rossicus from Berezhekovo, IAE 268. Occlusal view. most biostratigraphically important in the and hydromorphic fossil soils. These depo- non-glacial zone of Northern Asia. Thick sits mainly have lacustrine-alluvial gene- Quaternary sediments (seven alternated sis, fill the pits in lower layers and reach sub aerial and sub aquatic formations) the thickness of 45 m. In Kuznetsk Basin, with a great number of fossil bone remains Kedrovka Formation is associated with uncovered in opencast coalmines. More the most of Pleistocene large mammals than 60 taxa of the carnivores, probosci- remains. In the morphology and ecologi- des, perissodactyls and artiodactyls of cal peculiarities, recorded species from various geological age were recorded here, lower and upper levels of this formation allowing reconstruction of the faunal his- belong to the different faunistic comple- tory in the south of Western Siberia from xes. the Early Pleistocene to Holocene, to trace In the basal part, represented by the phylogenetic lines in basic mammalian Krasnogorsk Member, the fossil remains groups, and also to elucidate the successi- contain Gulo cf. schlosseri, Mammuthus tro - ve stages of their evolution gontherii (corresponding in its morphome- (FORONOVA, 1982, 1986, 1998, 1999, trical data to those of this species from in press). Süßenborn locality in Germany), Equus The bear remains (mandibles and pos- mosbachensis, Rangifer sp., and a very large tcranial elements) have been recorded in bison, Bison aff. priscus. This fauna is corre- the Kuznetsk Basin in situ at different lated with the Tiraspolian and Viatkian stratigraphical levels. The mandible IG faunas in East Europe and West Siberia 328 of Ursus rossicus was found in and the Cromerian faunas of We s t e r n Latyshovo levels of Kedrovka Formation Europe. in Bachatsk quarry. Kedrovka Formation The overlying Latyshevo Member con- represents bluish-gray loam and clay, plas- tains Ursus rossicus, Panthera spelaea, tic, flaky, with wood debris, basal shingle Mammuthus aff. chosaricus, Equus aff. tauba - 378 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001) chensis, E. ex gr. mosbachensis-germanicus, the riverbank. These were not associated Coelodonta antiquitatis, Cervus elaphus, with geological layers, but mainly origi- Megaloceros giganteus, Bison priscus. The nated from the Middle and Late teeth of M. aff. chosaricus, in comparison Pleistocene levels. The degree of conserva- with its ancestor M. trogontherii, differ in tion of the bones allows suggesting that less enamel thickness, the length of plates the main part of the bone assemblage had and larger plate frequency.