Cadernos Lab. Xeolóxico de Laxe ISSN: 0213-4497 Coruña. 2001. Vol. 26, pp. 373-398

Pleistocene small cave ( rossicus) from the South , 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 inhabited the steppe regions in early Middle and Late . By odontological characters it is more close to U. r. rossicus from Krasnodar, than to U. rossicus uralensis from Kizel Cave in . Discriminant analysis, based on measurements of lower cheek teeth of the cave 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 Ursus rossicus Borissiak, findings are also associated with the zones 1930 has been widely distributed in of spreading of the Pleistocene grass com- Northern 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 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 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 River (Omsk Region), River (Krasnyi Yar), (Strashnaya Cave, Denisova MATERIAL AND METHODS Cave), (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 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 , 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 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 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. Generally, the been formed in the Later Pleistocene. species composition and evolutionary level Castor fiber, Canis cf. lupus, Ursus cf. denin - of the fauna representatives correspond to geri (large form), Crocuta spelaea, Panthera the Holstein, which conform to spelaea, Equus caballus subsp., E. cf. prze - interglacial in Western Siberia. walskii, E. hemionus, Dicerorehinus kirchber - The mandible IG 10128 was found in g e n s i s, Coelodonta antiqutatis, M e g a l o c e r o s the sediments of Krasnobrodsk Formation giganteus, Cervus elaphus, Alces alces, Bison in Mokhovo quarry. These are lacustrine- priscus, Saiga borealis were recorded here alluvial loam with ferruginous shingle at (ALEXEEVA, 1980). its base and cryoturbations and frost- Kurtak archaeological region is situa- wedge casts in its upper part. ted in the south of Middle Siberia, in the Accompanied fauna is represented by Northern Minusinsk Basin. The sites were remains of Vulpes vulpes, Panthera spelaea, discovered after the flooding of the Yenisei Equus przewalskii, Equus ex gr. gallicus, River valley and forming of Krasnoyarsk Equus aff. hydruntinus, Coelodonta antiquita - Reservoir. The thickest Quaternary sedi- tis, Cervus elaphus, Megaloceros giganteus, ments of various origins are exposed in the Alces alces, Rangifer tarandus, Bison priscus, left riverbank, at Berezhekovo sites. There S a i g a sp., and Mammuthus primigenius. are opened here the ancient alluvial depo- Both thin and thick-enamel variations of sits of high Yenisei terraces, colluvial for- mammoth teeth can be distinguished. The mations (filling of the former ravines cut- first group may correspond to the begin- ting through terraces), and covering, ning of the Weichselian, while the second mainly eolian loess-like loams and sandy one having the radiocarbon dates 39.1, loams, containing several paleosoil hori- 31.9 and 28.9 thousand years belongs to zons. At the present time these sections the Middle Weichselian (Karginsk) war- have been examined geologically, archaeo- ming. Creating of these formations finis- logically, and paleontologically, and ther- hes apparently in the Late Weichselian moluminescent and radiocarbon data have cooling, as evidenced by cryoturbation been obtained (DROZDOV, et al. 1990a, features in the uppermost part of the 1992; DROZDOV, et al. 1990b). The sequence. bear remains described in this study, have The locality Krasnyi Yar that is the been found at the Berezhekovo 2,3 and 4 place of finding the skull ZIN 32748 and sites in the Middle and Late Quaternary mandible ZIN 35075, is situated 0 t Ob eroded brown loams, in the basement of River in Tomsk Province. A lot of bones section, and on the river beach. There have been gathered over several years on were found remains (determineted by CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 379

I.Foronova) of Canidae gen. indet., arch is thickened; its height equal to 40 H o m o t h e r i u m sp., Panthera spelaea, mm. The infraorbital foramen is small, Mammuthus primigenius, Mammuthus sp., situated over the posterior part of 1. Equus sp. (large form), Equus aff. hydrunti - Nasal bones are short, narrow, in anterior nus, Equus cf. przewalskii, Coelodonta anti - half being parallel to the palate. Their quitatis, Cervus elaphus, Megaloceros gigan - anterior end is finished at the level of 4. teus, Alces alces, Rangifer tarandus, Capreolus Nasal aperture is large, nearly as high as capreolus, Ovis cf. ammon, Bos sp., Bison pris - broad. The palate is comparatively broad. cus. The mandible IAE 36 was recorded in Its breadth increases greatly in front of the Berezhekovo 2, IAE 1681 in Berezhekovo anterior border of 4 (68.3 mm) to the 4, IAE 93, 330, 331, 455 in Berezhekovo canine alveoli (89 mm). The distance bet- 3-4. These localities are also associated ween medial margins of the latter exceeds with the Middle and Late Pleistocene that between 2, resembling in this featu- small mammals fauna (KRUKOVER & re the specimens from Krasnodar CHEKHA, 1999). (BORISSIAK, 1932). The length of inci- sors row is 62.4 mm. The anterior premo- DESCRIPTION lars P1-3 are absent; ALEXEEVA (1980) mistakenly considered the alveolus of P3 as an anterior alveolus of P4. The length of Skull post canine diastema is 38.5 mm. The dis- tance from the level of P4 anterior margin The skull of the bear from Krasnyi Yar to I1 alveolus is somewhat shorter than (ZIN 32748) has only the facial portion. the length of P4-M2. Cheek teeth are heavily worn. Their occlu- This specimen is peculiar in a rather sal surface nearly lost enamel, demonstra- posterior position of incisive foramina, ting the patches of dentine. Small size and which exceed beyond the level of posterior slender canines (length at enamel border edges of canines. This is not characteristic 16.3 mm, width 12.7 mm) indicate this for the cave bears and is probably caused specimen as an old female. by the very old age of the individual. Frontal bones are convex, abruptly rai- The skull ZIN 32748 in its size and sed over the nasal bones. The postorbital morphology is similar to those of U. rossi - process is short but broad and blunt. c u s from Krasnodar and Kizel Cave Orbits are placed closer than minimum (BORISSIAK, 1932, VERESHCHAGIN breath between orbits in the nominative & BARYSHNIKOV, in press). subspecies from Krasnodar (BORISSIAK, Measurements (mm): "snout" length - 1932). The orbit is comparatively large 138, length C1-M2 - 137.5, length P4- (greatest diameter 52 mm) and directed in M2 - 79.6, least breadth between the its upper portion more anterior than in U. orbits - 74.6, greatest palatal breadth - 84, arctos (ALEXEEVA, 1980). There are two breadth at the canine alveoli - 89, breadth lacrimal foramina approximately equal in between for. infraorbitale - 73.8, greatest size. The anterior root of the zygomatic breath of nasal opening - 57.6. 380 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001)

The skull from Berezhekovo is com- size common to U. rossicus. Anterior pre- pressed and belongs to the young indivi- molars p1-p3 are absent, the length of dual. The palatine between posterior diastema changes from 34.5 to 46 mm. molars is narrow. The anterior premolars The body of the mandible is high; the line are absent. of its inferior border is curved. The height of the body is considerably decreasing Mandible anteriorly, reaching its minimum at the diastema. The angular process rising, the Most specimens have worn to heavily condylar process is situated at the level of worn cheek teeth. In Berezhekovo, only articular surface. There are two mental three of 19 mandible fragments studied foramina, the posterior one being larger may be attributed to young . The and situating below p4, and sometimes it specimen from Mokhovo quarry also is divided in several small openings. belonged to a young bear. By the total size and the length of the Males of U. rossicus were probably dental row, the samples from Siberia are much larger than females very similar to those of U. rossicus from (VERESHCHAGIN & BARYSHNIKOV, Krasnodar (BORISSIAK, 1932) and from in press), sexual dimorphism is shown Kizel Cave (table 3). They are somewhat mainly in the canine size. In the material smaller on average than mandibles of U. investigated, three mandibles were attri- savini from Forest Bed, but this difference buted to males due to their greater total is not statistically reliable. length and width of the lower canine Factor analysis has been carried out for (table 1, 2). 11 measurements of male mandibles. The Mandibles demonstrate structure and results are shown in the figures 5, 6 and in

Table 1. Sizes of mandibles in males of Ursus rossicus from the South Siberia. CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 381

Table 2. Sizes of mandibles in females of Ursus rossicus from the South Siberia.

Table 3. Comparison of mandibles in Ursus rossicus.

table 4. In plote of the Factor 1 (c1-m3 morphometrical resemblance not only length, m2 length, m2 width) and Factor between the U. rossicus from Siberia and 2 (m1 length), the samples are divided in from Ural, but also the similarity of this two groups. The first one comprises U. small cave bear with U. savini. savini from Forest Bed and U. deningeri from Kudaro 1 Cave; the second contains Dentition U. rossicus from Siberia localities and from Kizel Cave (figure 5). In space of the Upper teeth. Upper cheek teeth of Factor 2 and Factor 3 (p4-m3 length, m1 the skull ZIN 32748 are heavily worn and width), the sample of U. deningeri appea- are not suitable for a morphological analy- red to be separated from others (figure 6). sis. Therefore only M2 has been measured Despite the fact that we dealt with a in this specimen. Additionally, three sam- rather small collection, we can suggest a ples of this tooth with obliterated occlusal 382 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001)

Figure 5. Plot of factor scores of Factor 1 and Factor 2 from principal component analysis of man- dibles. d- , Kudaro 1 Cave; f- U. savini, Bacton Forest Bed; k- U. rossicus, South Siberia; z- U. rossicus, Kizel Cave.

Figure 6. Plot of factor scores of Factor 2 and Factor 3 from principal component analysis of man- dibles. The symbols are as in figure 4. surface were recorded in Berezhekovo. On (table 5). The paracone is large, with a a Berezhekovo skull, judging from photo- blunt top. One specimen demonstrates the graph, the upper molars bear numerous lack of the additional tubercle in front of additional tubercles. the paracone, which is represented on the M2. The greatest length and width of teeth from Krasnodar (B O R I S S I A K, this tooth is correspondent to those in U. 1932) and is commonly present in U. spe - rossicus from Krasnodar and Kizel Cave laeus. The metacone is undivided, conside- CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 383

Table 4. Correlations of characters with the first three principal components for mandibles.

Table 5. Comparison of cheek teeth in Ursus rossicus. rably smaller than the paracone, and its Lower teeth. The lower cheek teeth apex is moved anteriorly. The row of lin- are more numerous; all of them were retai- gual tubercles in all specimens is heavily ned in mandibles and show the various worn. In U. savini, this tooth is on avera- degrees of crown abrasion. Relative values ge somewhat shorter and narrower, and in of the average length for p4, m1, m2 and U. deningeri from Westbury, it is, on the m3 are 16,3-27,7-29,1-26,9%. For this contrary, longer and wider, but in both index, the small cave bear from Siberia is cases the differences are not statistically similar to U. rossicus from Kizel Cave and reliable. It differs more noticeably with Krasnodar, although the former has relati- that of M2 in U. spelaeus from Odessa, vely longer m1 (29,1%). All the cave where it is apparently larg e r bears show m1 on average is shorter than (BARYSHNIKOV, 1998). m2, with exception for U. savini, having 384 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001) in contrary m1 longer than m2 (a primiti- cus, Siberia (n=5), U. spelaeus from Arcy- ve character inherited from U. etruscus). sur-Cure (n=41), from Odessa (n=40), and p4. The crown is of the equal width in from Secrets Cave (n=13). The results of the anterior and posterior parts. The pro- this analysis are present in figure 9, which toconid is high, blunt-pointed, and shows the centroid of each group plotted without crests. Its base bears two large onto the first two canonical variates. A lit- adjacent cusps on the lingual side (paraco- tle over 85% of dental variation is explai- nid and metaconid), these sometimes ned by the first canonical axis, 13% by the being linked with a small denticle (figure second. The first canonical variate discri- 7, 8). The paraconid commonly extends in minates greatest width, on the second front of the protoconid. A slender ridge canonical variate including greatest runs from the metaconid posteriorly. The length (table 6). The bivariate plot of cen- posterior tooth part is abruptly bent back- troids clearly shows an ordination into 2 ward, without additional tubercles, which separate groups, with savini/deningeri are recorded in the Krasnodar specimens samples on the one hand and all the other (BORISSIAK, 1932). There are two well- samples on the other. Squared separated roots. Mahalanobis distances between members The multiple discriminant function of these groups are high (from 5.59 to analysis based on three measurements 14.5). In the second group, which con- (greatest length, greatest width, and dis- tains U. rossicus from Siberia, the distances tance between paraconid and metaconid are lower (less than 3.87). The Siberian tips) was carried our for six samples: U. sample shows statistically reliable diffe- savini (n=10), U. deningeri (n=8), U. rossi - rences only from U. deningeri (p<0.001).

Figure 7. Lower premolar p4 of Ursus rossicus from Berezhekovo. Occlusal view. Figure 7. Plot of centroids for each locality on the plane canonical variate 1 and 2 for p4. CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 385

Figure 8. Lower cheek teeth p4 (a-d) and m1 (e- g) of Ursus rossicus from South Siberia. Lingual view. a-c, e-f - Berezhekovo, d- Bachatsk, g- Mokhovo.

Figure 9. Plot of centroids for each locality on the plane canonical variate 1 and 2 for p4.

Table 6. Correlations of canonical variate for p4. 386 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001)

Based on the results of this analysis, we hypoconid is large, with well-developed may conclude that p4 proportions in U. lingual part. The entoconid is in the form rossicus are similar to those of U. spelaeus, of a longitudinal ridge consisting of two being different with those U. savini, and three closely placed cusps; their length more sufficiently from U. deningeri. gradually increases backwards (figure 8). Among teeth of U. rossicus studied, the At the place of trigonid and talonid con- mean of the greatest length is at a maxi- tact, there is a weak labial cingulid. mum in Krasnodar and minimum in The discriminant analysis based on Kizel Cave (table 5), and the differences seven measurements (greatest length, between them are reliable for this measu- length of trigonid, length of posterior rement (p<0.01). The greatest crown entoconid tubercule, length of anterior width in the all samples slightly varies. entoconid tubercule, width of trigonid, U. etruscus and U. maritimus exhibit a width of talonid and width in the middle simple p4 without additional cusps. U. part of crown) was carried out for seven a r c t o s sometimes demonstrates a weak samples: U . s a v i n i (n=7), U. deningeri paraconid. Cave bears are characterized in (n=11), U. rossicus from Siberia (n=6) and the complication of the crown with acces- from Kizel Cave (n=12), U. spelaeus from sory tubercles ("molarization"). In U . A r c y - s u r-Cure (n=31), from Odessa deningeri, there is a paraconid and occasio- (n=29), and from Secrets Cave (n=8). The nally a small metaconid, these cusps being results of this analysis are present in figu- usual for U. savini. Both species show the re 10. Nearly 86% of dental variation is two cusps well separated. In U. rossicus, explained by the first canonical axis, paraconid and metaconid are larger and almost 10% by the second. The first cano- more approached together; sometimes nical variate discriminates greatest length additional tubercles are developed near and length of posterior entoconid tubercu- paraconid and metaconid and on the pos- le, on the second axis length of anterior terior crown side. This trend is continued entoconid tubercule contributes to discri- in U. spelaeus, demonstrating the larger mination (table 7). The plot of centroids tooth size and dilation of the posterior demonstrates division into 3 separate tooth part; the paraconid is slightly groups: 1) U. savini and U. rossicus, 2) U. moved anteriorly and together with the d e n i n g e r i and 3) U. spelaeus. Squared metaconid is separated from protoconid Mahalanobis distances between the first by a deep groove. Consequently, p4 in U. and the second groups (from 5.73 to 6.42) rossicus exhibits an advanced morphology are noticeably lower than between the (tooth enlargening, development of the second and third ones (from 7.99 to additional crown structures). 11.38). In the first group, the both sam- m 1. Lower carnassial tooth in the ples of U. rossicus are well-separated from Siberian samples is considerably wide. The one another (Mahalanobis distance 9.17), protoconid is considerably lower than the the Siberian specimens being closer to U. metaconid. The latter is moved posteriorly savini (1.17). Statistically reliable differen- and divided in two or three denticles. The ces (p<0.001) are observed between all CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 387

Figure 10. Plot of centroids for each locality on the plane canonical variate 1 and 2 for m1.

Table 7. Correlations of canonical variate for m1. samples with the exception of U. rossicus deningeri, this place is approximately as from Siberia/U. savini. Thus the m1 pro- wide as trigonid. U. etruscus, U. maritimus portions among U. rossicus are rather diffe- and U. arctos show the complicated meta- rent, but demonstrate more resemblance conid and entoconid, the latter consisting with those in U. savini, than with other of one or two cusps. Cave bears demons- cave bears (in the least degree with U. spe - trate a trend toward further development laeus). of entoconid portion. In U. rossicus, U. In the samples of U. rossicus, the lon- savini and U. deningeri, it is formed by two gest m1 has been recorded in Kizel Cave or three adjacent tubercles, while in U. (table 5), however its differences in this spelaeus the entoconid more often has a measurement with Siberian and shape of two large well-separated tuber- Krasnodar specimens are not statistically cles of almost equal height. Thus, U. rossi - reliable. cus demonstrates a rather primitive m1 In U. spelaeus, the crown is most structure. narrow at the trigonid and talonid junc- m2. This tooth is rectangular in shape, tion but in U. rossicus, U. savini and U. with talonid wider than trigonid. The 388 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001) protoconid is undivided, situating below of U. savini (5.76, 5.81). Statistically relia- the metaconid. The latter is divided in ble distances (p<0.001) seperate all sam- two or three nearly equal cusps, the mid- ples, with the exception of the samples dle one being the longest. This tubercle is from Siberia and Kizel Cave. The results not moved medially, which often takes indicate the resemblance of both samples place in U. spelaeus. The hypoconid is of U. rossicus in m2 measurements, but wider than the entoconid. The latter is they are well separated from the other longer than hypoconid and consists of two collections examined. Among cave bear l a rge and well-separated tubercles. species, the proportions of m2 in U. rossi - Sometimes an additional denticle between cus are similar to those of U. savini, being the metaconid and the entoconid occurs. more remote from U. deningeri, and espe- The discriminant analysis based on cially from U. spelaeus. The morphometri- seven measurements (greatest length, cal resemblance of U. rossicus and U. savini labial length of trigonid, lingual length of in this character has been already noted by trigonid, labial length of talonid, lingual KURTÉN (1959). length of talonid, width of trigonid, and Three samples of U. rossicus are similar width of talonid) was carried out for seven in m2 length (table 5), the collections samples: U. savini (n=11), U. deningeri from Krasnodar and Kizel Cave being sig- (n=31), U. rossicus from Siberia (n=9) and nificantly distinguished in the greatest from Kizel Cave (n=13), U. spelaeus from width of crown (<0.02). A r c y - s u r-Cure (n=43), from Odessa The m2 has a complicated metaconid (n=37), and from Secrets Cave (n=8). The and the large cusp of entoconid in U. etrus - results of this analysis are presented in cus, U. maritimus and U. arctos but the lat- figure 11. More than 76% of dental varia- ter is associated anteriorly with two or tion is explained by the first canonical three small denticles. In contrary, U. spe - axis, 11% by the second. The first canoni- laeus demonstrates two-cusped entoconid cal variate discriminates labial length of complicated by additional enamel folds. talonid, greatest length and width of tri- There is a cingulid on the crown lingual gonid, on the second axis lingual length of side, which is only slightly developed in talonid contributes to discrimination U. rossicus. (table 8). The plot of centroids demonstra- m3. The m3 crown is somewhat elon- te an ordination into 3 separate groups: 1) gated, with straight anterior and round U. savini and U. rossicus, 2) U. deningeri, posterior edges. It is shorter than that of and 3) U. spelaeus. Squared Mahalanobis m1, and wider than m2 crown. The large distances between the first and second talonid bears a clear notch on the labial groups (from 2.95 to 6.74) are approxima- tooth side. The protoconid is longer than tely the same to those between the second metaconid. The latter is high and divided and third ones (from 4.57 to 8.94). In the in two tips. The hypoconid is large, appro- first group, both samples of U. rossicus aching the protoconid. The entoconid is situate more close with one another represented by a serrate ridge. The crown (Mahalanobis distance 1.75), than to that area between tubercles is large and rugous. CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 389

Figure 11. Plot of centroids for each locality on the plane canonical variate 1 and 2 for m2.

Table 8. Correlations of canonical variate for m2.

The discriminant analysis based on spelaeus on the one hand and all the other four measurements (greatest length, examined samples on the other. Squared length of talonid, greatest width, and Mahalanobis distances between samples of width of talonid) was carried out for six these groups are rather low (from 2.59 to samples: U. savini (n=11), U. deningeri 5.06). In the first group, all the samples (n=47), U. rossicus, Siberia (n=6), U. spe - distribute very close to one another laeus from Arcy-sur-Cure (n=41), from (Mahalanobis distances from 0.16 to Odessa (n=42), and from Secrets Cave 1.65). Differences between the samples of (n=8). The results of this analysis are pre- U. rossicus and U. spelaeus are statistically sented in figure 12. Nearly 74% of dental reliable (p<0.001); there are no consistent variation is explained by the first canoni- differences between the samples of the cal axis, almost 16% by the second. The first group. Based on the results of this first canonical variate discriminates inclu- analysis we may conclude that by m3 des all measurements (table 9). The plot of measurements U. rossicus is more similar centroids clearly shows an ordination into to U. savini/deningeri, than to U. spelaeus. 2 separate groups, with all samples of U. Although the Krasnodar collection 390 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001)

Figure 12. Plot of centroids for each locality on the plane canonical variate 1 and 2 for m3.

Table 9. Correlations of canonical variate for m3. demonstrates the largest m3 in three sam- General remarks ples examined of U. rossicus (table 5), only the specimens from Krasnodar and Kizel Comparison of the lower cheek teeth Cave reliably differ in the greatest crown size in U. rossicus from Siberia, Krasnodar width (P<0.02). and Kizel Cave indicates a large metric In U. etruscus, U. maritimus and U. arc - resemblance of the first two samples. In tos, m3 is moderate in size, resembling an contrast, there are reliable differences elongated triangle or oval in occlusal view. (p<0.01) between the material from Commonly it is not clearly divided into Siberia and Kizel Cave by the greatest trigonid and talonid. In contrary, the cave width of p4, and between samples from bears and especially U. spelaeus and U. ros - Krasnodar and Kizel Cave by the greatest sicus are characterized by an increase of length of p4, width of talonid in m2 and tooth size, mainly of its talonid being well the greatest width of m3. Cluster analysis separated from trigonid. based on the average length and width of the lower teeth p4, m1, m2 and m3, has also confirmed (figure 13) that the sam- CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 391

Figure 13. Similarity dendrogram of samples of genus Ursus based on length and width means of lower cheek teeth p4, m1, m2 and m3. ples from Siberia and Krasnodar constitu- VERESHCHAGIN & BARYSHNIKOV te one cluster, while the sample from in press). The specimens examined resem- Kizel Cave groups with that of U. savini. ble in size the corresponding bones from The obtained results of the discrimi- Kizel Cave (table 10). The single fourth nant analysis show the resemblance of U. metacarpal, unlike those from Kizel Cave, rossicus with U. savini in proportions of the does not have the incision on the distal lower molars m1, m2 and m3. However in articulate ridge. proportions of the lower premolar p4, the former species is more close to U. spelaeus. CLADISTIC ANALYSIS OF CAVE The enlargement and complication of p4 BEARS may be treated as a specialization of U. ros - sicus to consume rough herbaceous food. The subgenus Speleartos E.Geoffroy, 1833 includes four species: U. savini Postcranial skeleton Andrews, 1922 (early Middle Pleistocene), U. deningeri von Reichenau, The isolated limb bones of a small 1904 (Middle and Late Pleistocene), U. cave bear have been found in Mokhovo rossicus Borissiak, 1930 (Middle and Late quarry and in Berezhekovo. To identify Pleistocene) and U. spelaeus Rosenmüller, their sex we used the data on the size sex 1794 (late Middle and Late Pleistocene). dimorphism in U. rossicus, established for The first and fourth species are found in the sample from Kizel Cave (see Europe only, the both others are found in 392 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001)

Table 10. Sizes of limb bones in Ursus rossicus from South Siberia. the many localities of the Northern 4. Body of the mandible: relatively Eurasia. low, its inferior border being straight (0), We scored seven ursid taxa and 17 cha- high, and the inferior border is curved, racters of skull, cheek teeth, limb bones, forming a prominence under m2 (1). and body size (see table 00). We used U. 5. Entepicondylar foramen of humerus: etruscus G.Cuvier, 1823 as an outgroup. present (0), absent (1). Among recent This species is considered to be ancestral bears this foramen is observed in primiti- both for the cave bears and the recent spe- ve and , and is absent cies of the genus Ursus (U. arctos and U. in the representatives of all other genera. m a r i t i m u s) (E R D B R I N K, 1953, In U. etruscus, this feature varies (MAZZA KURTÉN, 1968). & RUSTIONI, 1992). Characters. 1. Frontal profile: gentle 6. Metacarpal and metatarsal bones: (0), steep, so the frontal bones are strongly short (0), relatively long (1). raised over the nasal bones (1). 7. Ridge of the distal articulate surface 2. Nasal bones: long, terminating on metacarpal and metatarsal bones: anteriorly over the posterior edge of the round, without incision (0), with small upper canine (0), short, the anterior end incision on the distal border (1). situated over 4 (1). 8. Premolars P2-3/p2-3: present com- 3. Condylar process: lies at the level pletely (0), present partly (1), absent (2). with the occlusal surface of the lower 9. Anterior premolars P1/p1: present cheek teeth (0), elevated higher than the (0), absent (1). level of the latter (1). 10. Protocone of P4: simple (0), subdi- CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 393 vided or associated with additional tuber- confirm the distinct taxonomic status of cles (1). this subgenus. Peculiarity of the cave 11. Medial wall of the metacone P4: bears is caused by the high rate of their without tubercle and enamel fold (0), morphological evolution, and mainly by those present (1). changes of their masticatory system, pro- 12. Talon of M2: short and relatively voked by transition from omnivorous to narrow (0), long and moderately wide (1), chiefly vegetable diet. long and very wide, its inner field with Relationships within S p e l e a r c t o s a r e large rugae (2). The genus Ursus is charac- determined by two synapomorphies (cha- terized by the elongation of 2 talon, racters 14(2) and 3). These indicate the which is secondary reduced in U. martimus further specialization of the cave bears to (MAZZA, et al. 1995). various plant food. 13. Lower carnassial tooth m1: large, The obtained phylogenetic tree gene- longer than m2 (0), slender, shorter than rally agrees with the data on the sequence m2 (1). of cave bear species appearance in the geo- 14. Lower premolar p4: simple (0), logical record. U. spelaeus, for example, complicated with labial cusps (paraconid seems to be not only the most advanced and metaconid) (1), complicated with but also the youngest among them. labial cusps and additional tubercles (2). 15. Endoconid of m1: usually with a DISCUSSION single tip (0), with two tips or comb-sha- ped (1). Our investigation confirms that U. ros - 16. Talonid of m3: not developed (0), sicus was widespread in South Siberia in wide, separated from trigonid (1). Middle and Late Pleistocene. In propor- 17. Body size: medium or small (0), tions of the cheek teeth, the Siberian bear large (1). is more similar to the nominative subspe- Results. The data matrix was run by cies than to U. rossicus uralensis. Therefore, heuristic search option of PAUP and one the range of the nominative taxon inclu- tree was produced (TL=23, CI=0.826, ded the steppe zone from Ukraine to the RI=0.852). A parsimony analysis resulted Northern Caucasus in the west to in the phylogenetic hypothesis shown in Kazakhstan and south of Middle Siberia in figure 14. the east. The specimens from the Middle Among the selected features of the and Late Pleistocene do not show suffi- group U r s u s a r c t o s / m a r i t i m u s p l u s cient differences, possibly the former are Spelearctos character 5 are here considered somewhat larger. equivocal, but character 8(1) is considered BORISSIAK (1932: 190) suggested U. an unambiguous synapomorphy. Both rossicus was a steppe race of the cave bear, peculiarities separate this group from pri- which had more progressive dental struc- mitive ursids. tures than in U. spelaeus from Europe. Monophyly of the subgenus Spelearctos However, archaic characters of its denti- is supported by 12 synapomorphies. These tion allow him to treat these bears as a 394 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001)

Figure 14. Proposed phylogenetic hypothesis for the genus Ursus according to either accele- rated (ACCTRAN) optimization (L=23, CI=0.826, RI=0.852). See text for the descrip- tion of inferred synapomorphies 1-17. Character matrix is in table 11. Symbols: *- parallelism, R- revers.

Table 11. Character matrix for genus Ursus. parallel branch, originating from the com- level of differentiation of its dentition mon ancestor. does not exceed that in U. savini/deningeri. The obtained results partly support the Judging by material examined, publis- Borissiak opinion. U. rossicus has a specia- hed data (B O R I S S I A K , 1 9 3 2 , lized lower premolar p4. The trend to VERESHCHAGIN & BARYSHNIKOV, complication of this tooth among cave 2000), and distribution, we can imagine bears is demonstrating by U. savini (see that U. rossicus was a small bear of 1.5 m ANDREWS, 1922), reaches its greatest in length and 0.8 m in height in scapular development in U. rossicus and U. spelaeus. area. It seems to have been relatively slug- U. deningeri, in contrast, demonstrates gish and fed mainly on the vegetative simplification of p4. The morphology of parts of plants, roots, berries, fruits, and the other lower cheek teeth in U. rossicus is occasionally invertebrates, smaller verte- rather primitive and consequently, the brates, and carrion. This bear inhabited the open landscapes of plains and lower CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 395 mountains, where its small height allowed reconstruct for the cave bears, suggests it to hide from enemies in grass and relief their wide distribution in Eurasia and cavities. For over wintering, it evidently rapid adaptive radiation. We agree that dug earth dens at slopes and ravines, using they originated in the end of the Early caves rather seldom. Pleistocene. Among Spelearctos, a small U. The phylogenetic analysis carried out, savini seems to be the most ancient, retai- demonstrated the systematic unity of the ning to a considerable degree the omnivo- cave bears. They may be treated either as a rous characters of dentition (the lower car- subgenus of Spelearctos within the genus nassial tooth being large; additional Ursus, or as a distinct genus as has been tubercles on molars are slightly develo- repeatedly proposed by different authors ped). It was probably a forest species (BORISSIAK, 1932; VERESHCHAGIN, mainly. Its remains are recorded only in 1973; BARYSHNIKOV, 1998). Such a Europe (England, Austria), but we believe point of view may be supported by the fact it may be found also in Asia. To this spe- that distinctions, which subdivide the sis- cies, probably, belongs also U. "etruscus" ter species-group (U. arctos and U. mariti - gombaszoegensis from Gombaszog, in denti- mus), are quite often considered to be the tion of which one may recognize the ten- generic ones (for example, ELLERMAN & dency to U. savini and in some degree to MORRISON-SCOTT, 1966; McKeNNA U. deningeri (KRETZOI, 1938). & BELL, 1997). In the early Middle Pleistocene, U. ros - Our phylogenetic hypothesis (figure sicus and U. deningeri appeared. The former 14) is topologicaly identical with the retained a small size and settled the phylogenetic tree for the genus Ursus pre- Asiatic steppes. It did not reach the skele- sented by MAZZA & RUSTIONI (1994). ton proportions of U. spelaeus, but its THESE authors interpreted the hypothe- cheek teeth were already partly specialized tical ancestors for (deningeri + spelaeus) + for consuming vegetable food. (artos + maritimus), and for arctos + mariti - U. deningeri, in contrast, was an inhabi- mus clades as belonging to "Ursus gr. arc - tant of the forest and forest-steppe lands- tos". The latter was originated, in their capes of Europe, Caucasus, Central Asia opinion, from the early Ursus aff. etruscus, and south of Siberia. Its size has increased. which was more omnivorous than the later The crowns of the upper and lower molars U. etruscus. Really, the retained became more complicated but even the in dental morphology many features, latest representatives of the species did not which are primitive for the cave bears. reach the level of complication observed in However, the hypothesis of the origina- U. spelaeus. tion of the both cave and polar bears from In the late Middle Pleistocene, the U. arctos, is unacceptable in our opinion, forms transitional to U. spelaeus had appe- because if this a case U. arctos would repre- ared in Europe, this species predomina- sent the paraphyletic association (see also ting in the Late Pleistocene faunas. These TALBOT & SHIELDS, 1996). were very large bears, inhabiting various The evolutionary scenario, which we plain and mountain biotopes. In compari- 396 BARYSHNIKOV & FORONOVA CAD. LAB. XEOL. LAXE 26 (2001) son with U. rossicus and U. deningeri, it David (Paris), Dr. A. Forsten and Dr. M. shows the further strengthening of masti- Fortelius (Helsinki) for the help in study catory structures, which were necessary for the collections. We are also obliged to more effective fragmentation of plant Prof. N. Drozdov and Dr. N. Ovodov food. This contributed to the successful (Krasnoyarsk), Dr. A. Averianov and Dr. competition of U. spelaeus with other N. Abramson (St. Petersburg), and Dr. E. omnivorous large mammals of the Late Alexeeva (Izhevsk) who contributed in Pleistocene and species penetration far to this investigation. Dr. Svetlana north, up to the Northern Ural Baryshnikova (St. Peterburg) assists us in (Medvezhiya Cave at 620 N; see Kuzmina the work with manuscript. Dr. I. Barnes 1971). (Oxford) improved the English version of the paper. Photographs were made by ACKNOWLEDGEMENTS P.Labezkyi (Novosibirsk). We thank Dr. Aurora Grandal d’Anglade (La Coruña) for We express our gratitude to Prof. N. the wonderful organization of the Cave Vereshchagin (St. Peterburg), Dr. A. Bear symposium and editing our manus- Lister and Dr. A. Currant (London), Dr. F. cript. CAD. LAB. XEOL. LAXE 26 (2001) Pleistocene small cave Bear 397

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