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12 Baryshnikov* Gennady Bary s h n i kov Zoological Institute, R A S , S t . Petersburg C h ro n o l ogical and ge og r aphical variability of Crocuta spelaea ( C a r n i vo r a , H yaenidae) fro m the Pleistocene of Russia B a r y s h n i k o v, G., 1999 - Chronological and geographical variability of C rocuta spelaea ( C a r n i v o r a , Hyaenidae) from the Pleistocene of Russia - in: Haynes, G., Klimowicz, J. & Reumer, J.W. F. (eds.) – MA M M O T H S A N D T H E MA M M O T H FA U N A: ST U D I E S O F A N EX T I N C T EC O S Y S T E M – DEINSEA 6: 155-174 [ISSN 0923-9308]. Published 17 May 1999. Geographic variation in C rocuta spelaea dentition, beginning from the Middle Pleistocene, can be seen as specialization in western and eastern Eurasia. The sizes of C. spelaea increase from the south to the northwest and northeast. The hyena of the Primorski Krai had the largest teeth. C h ronologische en geografische variatie in Crocuta spelaea (Carnivora, Hyaenidae) uit het Russische P l e i s t o c e e n – Geografische variatie in het gebit van de grottenhyena, vanaf het Midden Pleistoceen, wordt beschouwd als een specialisatie in westelijk en oostelijk Eurazië. De maten van de grottenhyena nemen toe van het zuiden naar het noordwesten en noordoosten. De hyena van Primorski Krai had de grootste gebitselementen. Correspondence: Gennady Baryshnikov, Zoological Institute, Russian Academy of Sciences, 199034 St. Petersburg, Russia Keywords: C rocuta spelaea, Russia, geographic variability, Pleistocene I N T RO D U C T I O N The cave hyena C rocuta spelaea (GO L D F U S S , of bone assemblages of C. spelaea f r o m 1823) from Pleistocene Eurasia is usually European cave sites basically differs from regarded as an extinct northern subspecies of that in Africa, permitting us to assume diff e- the recent C. cro c u t a (ER X L E B E N , 1777) rences in behavior between C. spelaea a n d (Kurtén 1956, Werdelin & Solounias 1991). C. cro c u t a (Brugal et al. 1997). The cave This assumption, based upon a formal simi- hyena was probably an active predator of larity of skeletal features of both forms, is in Pleistocene ungulates and may have lived in fact debatable. The view that C. spelaea is a l a rger groups with a complex hierarchical separate species is supported by the more s t r u c t u r e . carnivorous specialization of its dentition, as compared to C. cro c u t a (Baryshnikov 1995, An ancestor of C. spelaea was C. sivalensis Baryshnikov & Averianov 1995). One also (FA L C O N E R E T CA U T L E Y, 1868) from the may assume that the cave hyena had adapta- Siwalik fauna of India, which during the tions to life in a climate that had a long Cromerian interglacial spread into the tempe- frosty period (adaptations predicted, for rate latitudes of Eurasia (Kurtén 1956, 1957). example, to be in features such as the deve- I refer to C. spelaea all Pleistocene discoveries lopment of a dense fur coat). The character of the genus C ro c u t a from Europe, western 155 MAMMOTHS AND THE MAMMOTH FAUNA DEINSEA 6, 1999 Figure 1 Localities with Pleistocene remains of Crocuta spelaea in Russia and adjacent terri t o ri e s : a - Cromeri a n - M i n d e l , b - Riss- W ü rm , c - W ü rm . 1 - Kazanka Rive r, 2 - Binagady, 3 - Golabiec, 4 - Prolom 2, 5 - Ilskaya I, 6 - Nasorog Cave, 7 - Smelov s k aya 2 C ave, 8 - A m a n - K u t a n , 9 - Kra s nyi Ya r, 1 0 - Bukhtarm i n s k aya Cave, 1 1 - Ust-Kanskaya Cave, 1 2 - Denisova Cave, 1 3 - Geographical Society Cave. Asia, middle Asia, Siberia and China; the on the localities of the last interglacial (Riss- genus evolved several subspecies replacing Würm) in Moldova (Vykhvatintsy), Ukraine each other in time and space (Kurtén 1956, (Iliinka, Kiik-Koba) and A z e r b a i j a n 1965, Bonifay 1971, Schutt 1971, Kurtén & (Binagady). We have a better knowledge of Poulianos 1977). the distribution of C. spelaea in the Early- Middle Würm (Fig. 1). In deposits of that In the former Soviet Union the earliest record period its abundant remains occur jointly of C. spelaea falls in the Cromerian/Mindel with a complex of Mousterian stone imple- transitional period. During that time interval ments. Cave hyena was found in Moldova, the remains of C ro c u t a are known from the Crimea, the Northern Caucasus, the Southern Ukraine (Opolie, Sinyakovo), Russian Plain Urals, middle Asia, Kazakhstan, and (Kazanka River), Northern Caucasus Southern Siberia eastwards up to the A n g a r a ( Treugolnaya Cave), and Kazakhstan River basin, while further in the east it appe- (Zhelesinskoe; Baryshnikov & Ve r e s h c h a g i n ars again in the southern Primorski Krai 1996). No significant discoveries are known where it penetrated from China (Baryshnikov for that area for the period from the Late & Vereshchagin 1996). In the Crimea and in Mindel up to the Riss, although remains of Middle Asia C. spelaea disappears by the C ro c u t a from Kyrgystan (Sel-Ungur Cave, beginning of the early Paleolithic, but in the Mindel or Mindel-Riss) and Moldova (Starye Urals and the Altai it could have survived D u r u i t o r y, Riss ?) could belong to the same until the main climatic minimum of the Late time interval. More information is available P l e i s t o c e n e . 156 BARYSHNIKOV: Pleistocene Crocuta from Russia During the Pleistocene European C. spelaea Russian Academy of Sciences, Novosibirsk demonstrated notable differences in the sizes (I A E); the Institute of A r c h a e o l o g y, Ukrainian of skull and teeth, observed when the disco- Academy of Sciences, Kiev (I A U ); the veries from deposits of glacial and interg l a- Institute of Systematics and Evolution of cial epochs are compared (Kurtén & Animals, Krakow, Poland (I S E ); l'Université Poulianos 1977, Klein & Scott 1989). Claude Bernard Lyon 1, Villeurbanne, France H o w e v e r, the tendencies of geographic varia- (U C B); l'Université Pantheon-Sorbonne, Paris, tion at different temporal levels of the France (U P S ); and the Laboratoire de Pleistocene are not sufficiently studied. In Geologie du Quaternaire, Marseille-Luminy, this paper the major tendencies of spatial and France (L G Q ) (Table 1). temporal variation of C. spelaea are discus- sed for the former Soviet Union and We s t e r n R u s s i a Europe. Most attention has been concentra- KA Z A N K A RI V E R. Complete skull with lower jaw from ted on molars, the sizes and proportions of the Kazanka River in Tatarstan (G M M 191); Middle which varied depending not only on the size Pleistocene (Cromerian-Mindel). of animals, but also on dietary specialization. IL S K AYA 1. Postcranial bones and lower cheek teeth (ZIN 19889). Mousterian open site at Il River (Kuban M AT E R I A L S Basin), Northern Caucasus; Early W ü r m . The materials studied include sixteen samp- NO S O R O G CAV E. Left mandible (ZIN 34477). A cave in les of fossil hyena remains from the collec- the South Urals; apparently dated to the Early W ü r m . tions of the Zoological Institute, Russian SM E L O V S K AYA 2 Cave. Approximately 45 C ro c u t a Academy of Sciences, St. Petersburg (Z I N ) ; remains (ZIN 34491).Upper Paleolithic site on Malyi the Geological and Mineralogical Museum of Kizyl River, South Urals. C ro c u t a remains come Kazan University, Kazan, Russia (G M M ); the from layer 5, lying beneath the cultural layer. Institute of Archaeology and Ethnography, Radiocarbon dates are: layer 3-4: 31,000 +/- 1500 y B P (GIN-8401); 41,000 +/- 1800 yBP ( G I N - 8 4 0 2 ) . KR A S N Y I YA R . Left mandible (ZIN 32752) from Ob River beach near village Krasnyi Yar in Tomsk Ta ble 1 The number of specimens studied. Region (Fig. 2). US T- K A N S K AYA CAV E. Postcranial elements and isolated teeth of hyena (ZIN 34404) within highly fragmen ted ungulate bones, in a Mousterian cave site on Charysh River in Altai; Karginian interglacial Mousterian deposits. DENISOVA CAVE. A few isolated bones and teeth (coll. IAE) from Mousterian (layers 12-20) and Upper Paleolithic (layers 9-11) site located on Anui River in Altai. Layer 21 has termoluminisciences dates 155,000 ± 31000 (RTL-546) and layer 14 as 69,000 ± 17000 (TL-9), and layer 11 radiocarbon date as >37,235 yBP ( S O A N - 2 5 0 4 ) . GEOGRAPHICAL SOCIETY CAVE. More than 170 bones and teeth, including fragmented skull (Fig. 3) and few lower jaws (ZIN 34478-34490) mainly from a depth of 80-140 cm below the surface (layer 4); Upper Paleo- lithic site located at Partizanskaya River (= Suchan River) in southern Primorski Krai.
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