Cadernos Lab. Xeolóxico de Laxe ISSN: 0213-4497 Coruña. 2001. Vol. 26, pp. 447-455

The updated record of cave bear and other members of the genus Ursus on the territory of Registro actualizado de Oso de las Cavernas y otras especies del género Ursus en la República Checa

WAGNER, J.

AB S T R A C T

During the 2nd half of 20th century, a succession of new Pleistocene localities was dis- covered and some classical paleontological sites were revised. From many of those pla- ces also bear remains were recorded. Nine localities provided large material of bear remains in good stratified layers. Bear record from Moravian localities were studied in details by Prof. R. MUSIL from . This paper presents a review of updated kno- wledge on this localities - especially about their stratigrafical position (based on micro- mammalian evidence) and basic information on bear remains studied.

Key words: Pleistocene, Ur s u s , cave bear, brown bear, Czech Republick, Bohemian karst, Moravian Karst Charles University. Faculty of Sciences. Department of Palaeontolog. Albertova 6.12843 Praha 2. CZECH REPUBLIC 448 WAGNER, J. CAD. LAB. XEOL. LAXE 26 (2001)

INTRODUCTION The most important informations are given by new discovered sites with both Palaeontological investigation of fossil large and small mammals together, which cenozoic mammals has a tradition in the allows morphometric studies of bears territory of the Czech Republic. Already combined with their exact stratification. from the begining of this investigations The most importent are as follows: remains of bears were among the most fre- 1) Holstejn (Moravian karst) quent finds. The first phase research star- Stratigraphy: zona B of Early Biharian st ted in area of Moravian karst in the 1 half 1 (FEJFAR & HORACEK, 1990) th of the 19 century (e. g. WANKEL made Stratigraphicaly significant taxa from in 1850 first skeleton of cave bear in the this locality: Microtus (Allophaimys) pliocae - Austria-Ungarn from bones founded in n i c u s KORMOS, Mimomys pusilus nd th Vupustek cave). From the 2 half of 19 (MÉHELY), Pliomys episcopalis MÉHELY, century lot of data about bear finds from P. lenki (HELLER), Hypolagus beremendensis (WANKEL. 1867; MASKA, (PETENYI) (according MUSIL, 1966). 1884, 1886; KNIES, 1891, 1893 etc.) and The others species of carnivores from from Bohemia (KREJCI, 1865; LAUBE, this locality: P a n t h e r a cf. g o b a s z o e g e n s i s 1874; WOLDRICH, 1890, 1893, etc.) (KRETZOI), Putorius cf. stromeri KOR- th has been recorded. At the end of 19 and MOS, Meles meles (LINNAEUS) (according st th during the 1 half of 20 century first MUSIL, 1966). lists of vertebrates fossils finds (incl. bears) Locality was discovered in 1965 were given from the area of Bohemia during limestone-mining. It was a chim- (WOLDRICH, 1897; KAFKA, 1901; ney cave infilled with terra rossa deposits. B AYER, 1905) and Moravia (KNIES, Fauna was published by MUSIL (1966). 1929; STEHLIK, 1929; SKUTIL & The locality provided about 30 items STEHLIK, 1932). The old authors deter- of bears, all identified as Ursus sp. by mined their finds mostly as U. spelaeus or MUSIL who compared them to the bears Ursus arctos/priscus. Some of these finds are of Biharian age (U. deningeri, U. mediterra - but lost, the most of remaining is not revi- neus, U. e. ? d. gombaszoegensis) but did not sed and without stratigraphical level or coidentified the Holstejn bears with anyo- nd st correct localization. Till in 2 half of 20 ne of them. The form was very near to U. century began modern palaeontological e. ? d. gombaszoegensis but exhibited some research. New and revised old localities more primitive features. are discovered, with emphasis to stratigra- The following items were published: phical signification. The most important fragment of mandible with M2 and M3, researchers of this period are FEJFAR in free C, fragment of mandible with C, and Bohemia (FEJFAR, 1956, 1961, 1993, a lot of fragments of long bones, hand and etc.) and MUSIL (MUSIL, 1959, 1962, food bones and ribs. 1965, 1968, etc.) in Moravia. The papers 2) Stranska Skala (Brno) of the last author brougt detailed studies Stratigraphy: Late Biharian 2 (FEJ- of cave and brown bear. FAR & HORACEK, 1990) CAD. LAB. XEOL. LAXE 26 (2001) The updated record of cave bear on Czech Republic 449

Stratigraphicaly significant taxa from Pitymys gregaloides HINTON, P i t y m y s this locality: Allocricetus bursae SCHAUB, s c h m i d t g e n i KRETZOI (according FEJ- Pliomys episcopalis M E H E LY, Mimomys FAR, 1961). savini HINTON, Pitymys hintoni KRET- The others species of carnivores from ZOI, Pitymys gregaloides HINTON (accor- this locality: Canis mosbachensis S O E R- ding LOZEK & FEJFAR, 1957 and GEL, Vulpes angustidens T H E N I U S , MUSIL & VALOCH, 1968) Mustela palerminea (PETENYI), Meles ata - The others species of carnivores from vus KORMOS, Xenocyon lycaonoides KRE- this locality: Canis mosbachensis S O E R- ETZOI, Hyaena brevirostris AY M A R D , GEL, ?Cuon sp., Vulpes angustidens THE- Panthera fossilis (VON REICHENAU), NIUS, Vulpes praeglacialis, ?Vu l p e s s p . , Panthera gombaszoegensis KRETZOI, Felis Xenocyon lycaonoides KREETZOI, Crocuta sp., Lynx issiodorensis (CROIZET & crocuta ssp., Hyaena brevirostris AYMARD, JOBERT), Homotherium moravicum (WOL- Gula schlosseri KORMOS, H o m o t h e r i u m DRICH) (according FEJFAR, 1961, moravicum (WOLDRICH), Lynx sp., Meles WOLSAN, 1993 and MAZUCH, 1997) atavus KORMOS, Panthera gombaszoegensis This is a karst filling - relict of an KRETZOI, Panthera pardus (LINNAEUS) extent cave entrance. It was studied by (according (MUSIL & VALOCH, 1968 FEJFAR (1956, 1961 etc.). From this and WOLSAN, 1993) filling exist detailed description of layers This is a classical palaeontologic site and their micromammalian remains, the- repeatedly investigated since the begin- refore all record of bears collected by ning of 20th Century. The history of inves- Fejfar is stratigraphically fixed. tigation was described in details by From this locality two bear species MUSIL (1965). Besides of the main sec- were described: U. deningeri and a smaller tion (site 4 by MUSIL), a number of diffe- form, which is usually reported as U. medi - rent fissures and caves nearby it was also terraneus (of course, it could be a member investigated. of Ursus gr. arctos sensu MAZZA & RUS- The finds of bears before 1945 was TIONY, 1994, in fact). The material is revised by MUSIL (1972) and new mate- rich. It is partly deposited in Charles rials was discribed by the same author University (FEJFAR collection), partly in (MUSIL & VALOCH, 1968; MUSIL, National Museum (PETRBOK collection) 1995). All Biharian bears were designated and a few pieces (about 14) in Museum of as U. deningeri. Bohemian karst in Beroun (LYSENKO 3) C 718 (Bohemian karst) collection). Stratigraphy: Late Biharian 2 (FEJFAR 4) Stranska Skala cave (Brno) & HORACEK, 1990); this is one glacial Stratigraphy: earlier part of Toringian cycle probably at the end of biharian. (FEJFAR & HORACEK, 1990) (based on Stratigraphicaly significant taxa from the micromammalian from cave clay) this locality: Allocricetus bursae SCHAUB, Stratigraphicaly significant taxa from Pliomys episcopalis MEHELY, Mimomys savi - this locality: Microtus gregalis type "grega - ni HINTON, Pitymys hintoni KRETZOI, loides", Microtus arvalis type "arvaloides", 450 WAGNER, J. CAD. LAB. XEOL. LAXE 26 (2001)

Dicrostonyx simplicior, Pliomys episcopalis The material is deposited in The National (according FEJFAR & HORACEK, 1990) Museum . It is a spacious cave where a large The Würmian (= Weichselian) finds number of bear remains was found just at are fairly poor. They were identified as the surface of deposits. These bears was Ursus arctos ssp. by MOSTECKY (1969). determinated as U. deningeri (MUSIL, SEI- Between these two bear-containing stratas TEL in litt. ex FEJFAR & HORACEK, is a considerable time gap from which no 1990). By MUSIL`s opinion (MUSIL in bear remains are available (W 1-2) (MOS- litt.) the bears remains are of the TECKY, 1969). Cromerian age (i.e. the late Biharian), the 6) Cave “Barova” (Moravian karst) microfauna obtained from the deposits by Stratigraphy: Upper Toringian (sensu HORACEK (in litt.) was apparently of FEJFAR & HORACEK, 1990); Würm the post-Biharian age. glacial ( MUSIL, 1959) 5) Chlupacova sluj (Bohemian karst) This is a typical bear-cave. It was dis- Stratigraphy: toringian (FEJFAR & covered in 1947. The section of the cave HORACEK, 1990); in this cave are the bearing a large concentration of cave bear interglacial and glacial deposits of Eemian bones was discovered in 1958 during the and Weichselian speleological investigation. The matere- I n t e rglacial micromammalian: rial was collected by R. MUSIL. Microtus arvalis, Apodemus sp., Glis sp. The most bear remains were in red- (FEJFAR in litt.) brownish (ferruginous) soil (W1-2). The others species of integlacialy car- Anyhow, the bones were found also in the nivores from this locality: Vu l p e s s p . , overlying layers but in much smaller com- Panthera spelaea (GOLDFUSS), Crocuta spe - centration. Excavated were only the W1-2 l a e a (GOLDFUSS) (according MOS- remains. Tooth (e. g. 6x P4, 17x M1, 22x 2 TECKY, 1969). M , 9x P4, 12x M1, 28x M2 and 18x M3) This is a remain of vertical karst cavity. from this layer were studied by MUSIL It was discovered during limestone (1959). The bears were determinated as mining at the end of 19th or beginning of Ursus spelaeus. 20th century. 7) Cave “Sveduv stul” (Moravian The R/W (= Eemian) interglacial bears karst) were studied by MOSTECKY (1963) and Stratigraphy: Upper Toringian (sensu determinated as Ursus arctos taubachensis. FEJFAR & HORACEK, 1990); Würm Besides the isolated tooth (e. g. 4x P4, 5x glacial (MUSIL, 1962) 1 2 M , 7x M , 3x M1, 2x M2 and 2x M3) and The others species of carnivores from postcranial elements (published by MOS- this locality: Panthera pard u s ( L I N- TECKY) a nearly complete skull is avaial- NAEUS), Crocuta spelaea (GOLDFUSS), ble from the interglacial layers of the site Canis lupus LINNAEUS, Vulpes vulpes (HORACEK in litt.). The most of the (LINNAEUS), Alopex lagopus seu corsac, items were collected in the sixties by Meles meles (LINNAEUS) (according J.Peterbok and latter by MOSTECKY. MUSIL, 1962). CAD. LAB. XEOL. LAXE 26 (2001) The updated record of cave bear on Czech Republic 451

This is a one of the typical palaeonto- Mustela cf. minuta (POMEL), Putorius cf. logical localities in Moravian karst. The putorius (LINNAEUS) (according MUSIL, investigation has begun here at the end of 1965) 19th century. This is a typical hyaena-cave. The site known since the end of 19th The results of the recent investigation century was newly studied during 1956- (1953-1955) was published by MUSIL 1958 by MUSIL (1965). (1962). The Pleistocene layers were subdivided The locality provided more than 700 into 5 sections (W1, W1-2, W2, W2-3, tooth and some bones (not so many). All W3). All these stratas were fossiliferous, were dated to the Würmian, covering and, in general, quite a large collection of either W1-2 (+? R/W), W2, W2-3 and bears was obtained from this locality. The W3. The bear remains were found in all most of remains belong to Ursus spelaeus, a the stratas. The largest number of remains few finds (remains of three mandubles) come from W1-2 (320 tooth) and W2 from W1-2 were designated as Ursus arctos (343 tooth). Both Ursus spelaeus and Ursus priscus. Teeth and bones were described in a r c t o s were determinated but most of details by MUSIL (1965) who compared remains belong to cave bear. tham with those of the cave bears from Worth of particular interest are some "Barova" cave and "Sveduv stul" and tooth from W1-2 and W2, called "tooth demonstrated morphometric changes with simple morphology". They are assigned during Pleistocene in morphometric pat- to cave bear, but have only main cusps, terns of cave bear populations and mor- simple inner field and are mostly without phometric differences between separate cingulum. These characters states are even populations of this species over such a here quite a rare, of course. small area as that of the Moravian karst. 8) Cave “Pod hradem” (Moravian The morphogenetic analysis of cave bears Karst) from cave "Pod hradem" was further Stratigraphy: Upper Toringian (sensu undertaken by G. RABEDER (1989). FEJFAR & HORACEK, 1990); Würm 9) Predmosti near Prerov glacial ( MUSIL, 1965) Stratigraphy: Upper Toringian (sensu Stratigraphicaly significant taxa from FEJFAR & HORACEK, 1990); würm this locality: Citellus citellus LINNAEUS, (MUSIL, 1965) Microtus gregalis (PALLAS), Microtus oecono - This locality ranks among the classical mus (PALLAS), Microtus nivalis (MARTIS) palaeontological and archaeological loca- ( according MUSIL, 1965) lity. The records come from an extensive The others species of carnivores from loess deposit very rich in fossil remains this locality: Panthera spelaea (GOLD- (the station of the Mammoth hunters). FUSS), Crocuta spelaea ( G O L D F U S S ) , The bear material was revised by MUSIL Canis lupus LINNAEUS, Vulpes vulpes (1964). (LINNAEUS), Alopex lagopus ( L I N- The former descriptions coidentified NAEUS), Martes sp., Mustela erminea LIN- the bear material with Ursus spelaeus. NAEUS, Mustela nivalis L I N N A E U S , During revision by MUSIL also a larger 452 WAGNER, J. CAD. LAB. XEOL. LAXE 26 (2001) collection of Ursus arctos priscus was here morphological and metrical comparision identified which come from the end of of this form with the cave bear from the W2 (or W2-3) or from the begining of cave "Pod hradem"., "Sveduv stul" and W3. MUSIL (1964) undertook a detailed "Barova".

A biostratigraphic table with stratigraphical position of refered localities (the base of the Biharian is a FAD of Microtus) (accordina FEJFAR et HORACEK, 1990) CAD. LAB. XEOL. LAXE 26 (2001) The updated record of cave bear on Czech Republic 453 454 WAGNER, J. CAD. LAB. XEOL. LAXE 26 (2001)

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