Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14:161–175, Wien 2005

Cave Bears (Ursidae, Mammalia) from the Trojuholník Cave ()

Martin SABOL*)

SABOL, M., 2005. Cave Bears (Ursidae, Mammalia) from the Trojuholník Cave (Slovakia). — Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14:161–175, Wien.

Abstract & HOLÚBEK, 1999) in a rock wall near northern end of the Prepadlé quarry (HOLEC et al., 1994) (fig. 2). The cave was Fossil remains of bears from the Trojuholník Cave in the named according to the shape of its triangular entrance Malé Karpaty Mts. have been analysed from the mor- (SILNICKÝ, 1949). Besides assemblage of fossil vertebrates phological, metric, and morphodynamic point of view. from the end of the Last Glacial and start of the Holocene They probably belong to the cave bear group (Ursus (HOLEC et al., 1994), the remains of cave bears have been spelaeus ROSENMÜLLER & HEINROTH, 1794) from the Mid- found here too (SILNICKÝ, 1949, 1950). dle Pleniglacial. Keywords: bears, Trojuholník Cave, Middle Plenigla- cial 2. Material and Methods

K. SILNICKÝ (1949) has collected the fossil remains of bears Zusammenfassung in 1948 during excavations in the Trojuholník Cave. They are deposited as a part of the fossil vertebrate collections of Fossile Reste der Bären von Trojuholník Höhle in den Klei- the Slovak National Museum - Museum of Natural History nen Karpaten sind aus morphologischem, metrischem und (SNM-MNH) in . From whole fossil material, 6 morphodynamischem Gesichtspunkt analysiert worden. isolated teeth, 7 skulls or their fragments and 15 mandibles Sie gehören wahrscheinlich zu der Höhlenbären Gruppe or their fragments were under study. (Ursus spelaeus ROSENMÜLLER & HEINROTH, 1794) aus dem Photos of fossils have been converted in the Corel-Photo Mittelpleniglacial. Paint 8 software. The measurements of studied ursid Schlüsselwörter: Bären, Trojuholník Cave, Mittelple- remains have been measured by standard method with niglacial callipers to 0.1 mm calibration precision. The standard deviation of measuring is 0.3 mm with 0.1 mm disper- sion and random error 4.2%. All measured data are in 1. Introduction millimetres. Terminology of main cusps on the tooth crowns is as fol- Fossil remains of mammals in cave deposits belong among lows - a designation ending with “-id” (e. g. protoconid) the most important records for the Quaternary stratigra- is used for cusps of lower teeth and ending with “-e” phy. Besides findings of micromammals, teeth and bones (e.g. protocone) for cusps of upper teeth. Ursid fossils of bears (especially cave bears) are the most frequently are classed to system according to MCKENNA & BELL found in caves. (1997). In the territory of Karst (Malé Karpaty Mts., Abbreviations: SNM-MNH (Slovak National Museum south-western Slovakia), some caves with records of – Museum of Natural History, Bratislava, Slovak Re- vertebrate fossils are situated (fig. 1). However, the most public). fossils have been found in the Trojuholník Cave. This cave is located near village of Borinka in the district. It is 35 m long, with opening 356 m above sea level (BELLA 3. Systematic Palaeontology

Class: Mammalia LINNAEUS, 1758 Order: Carnivora BOWDICH, 1821 ) * DR. MARTIN SABOL, Department of Geology and Pa- Suborder: Caniformia KRETZOI, 1943 leontology, Faculty of Science, Comenius University, Infraorder: Arctoidea FLOWER, 1869 Mlynska dolina, SK-842 15 Bratislava, Slovak Republic, Parvorder: Ursida TEDFORD, 1976 e-mail: [email protected] Superfamily: Ursoidea FISCHER DE WALDHEIM, 1817 162 Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14, Wien, 2005

0 1 2 3 km Figure 1: Location of the Tro- juholník Cave in the Borinka SpáleniskoSpalenisko Karst area (modified according 503 Volhovisko to HOLEC et al., 1994). 587 Z-796, Z-797, Z-802, Z-804, Z-805); 3 m3 dex. (Z-793, Stupavka Z-795, Z-800); 3 m3 sin. (Z- 792, Z-802, Z-805); 7 skulls TrojuholníkTrojuholnik Cave or their fragments (Z-452, Stupava Borinka Z-788, Z-790, Z-791, Z-794, Z-799, Z-801); 15 mandibles or their fragments (Z-789, Z-792, Z-793, Z-795, Z-796, Z-797, Z-798, Z-800, Z-802, Z-803, Z-804, Z-805, Z-806, Vydrica Z-807, Z-1001).

ZahorskaZáhorská BystricaBystrica Hrabovina 3.2. Description 434 The light to brown crowns of RacaRača teeth are mostly undamaged, LamacLamač but slightly worn to abraded. Vinohrady Some teeth as well as skulls DúbravkaDubravka and lower jaws are covered by fine-grained fluvial gravel in the most cases. Whereas two distinct lobes, separated by a conspicuous notch, are situated on the lingual side of I2 sin, a strong S L O V A K I A cingulum is developed on the Bratislava same side of I3 sin. with vis- BRATISLAVA ible crest extending medio- laterally from its top. One of five upper canines Family: Ursidae FISCHER DE WALDHEIM, 1817 has preserved a distinct distal crest. Two upper canines and two lower ones belonged Genus: Ursus LINNAEUS, 1758 probably to female individuals. The morphology of the both upper and lower premolars is specified by the classification of these teeth to single Ursus spelaeus ROSENMÜLLER et HEINROTH, 1794 morphotypes, determined by RABEDER (1983, 1992). On figs. 3, 4, tab. 1–4 the basis of that, it is possible to determine morphotypes A/D, E, and probably B for three P4s (two other P4s from damaged skull (Z-788) represent probably morphotype A 3.1. Material and A/D, but their exact determination is not possible for reason of their covering by a sediment) and C1, D1, and 1 I2 sin. (Z-452); 1 I3 sin. (Z-452); 2 C sup. dex. (Z-809, probably C3 for four p4s (tab. 5a-c). P4/4 standardised Z-810); 3 C sup. sin. (Z-452, Z-788, Z-812); 1 c inf. dex. index is 73.10 (or 62.03 in the case, when all P4s are used (Z-800); 3 c inf. sin. (Z-796, Z-797, Z-808); 1 canine for analysis). fragment (Z-811); 3 P4 dex. (Z-452, Z-788, Z-794); 2 P4 An indication of accessory cusp at the back side of para- sin. (Z-452, Z-788); 1 p4 dex. (Z-813); 3 p4 sin. (Z-789, cone is present on the crown of one from six M1s, whereas Z-792, Z-804); 4 M1 dex. (Z-452, Z-788, Z-794, Z-801); an indication of accessory cusp in front of metacone is 2 M1 sin. (Z-452, Z-788); 4 M2 dex. (Z-452, Z-788, Z- present on the crown of three M1s. Anyway, a distinct crest 794, Z-801); 3 M2 sin. (Z-452, Z-788, Z-801); 2 m1 dex. on the metacone inner side is developed in one case too. (Z-795, Z-800); 4 m1 sin. (Z-789, Z-792, Z-797, Z-802); 3 The both parastyle and metastyle were always developed. m2 dex. (Z-795, Z-798, Z-800); 7 m2 sin. (Z-789, Z-792, Two molars have a undivided protocone and a distinct SABOL, M., Cave Bears (Ursidae, Mammalia) from … 163

Figure 2: The Trojuholník Cave. S1-3 – pits (modified according 350.34 to HOLEC et al., 1994). notch between the protocone and paracone is present on S-3 one molar. The mesocone (metaconule), mostly smaller than hypocone, is divided to two cusps in one case. Similar to the protocone, two molars have a undivided hypocone. 350.34 The central area of M1s is either smooth or humpy. Only lingual cingulum is devel- oped. S-2 The paracone of two from six M2s has developed a 349.22 small accessory cusp at the back. Accessory cusps are also present either in front or at the back of metacone, which is same large as the paracone in one case. A blunt cusp (metastyle) is developed 348.90 behind the metacone on the crowns of four molars and 349.67 four crests forming X-like S-1 shape on the inner side of 351.34 metacone are present in one case. Three molars have the 354.30 protocone divided to two parts and two ones have this cusp divided to three parts. 355.36 The front part is mostly the largest. In most cases, the 5 m metaconule is undivided and entrance with an accessory cusp at the back of one tooth. A distinct crest consisting of two cusps is developed between the metacone and metaconule in one case. While the hypocone do not coincided with acces- sory cusps at the back crown margin, it has a small cusp developed at the back. The The paraconid of m2s mostly coincides with other cusps on talon of four molars is rounded and pointed in one case. the anterior crown margin and one molar has not developed The central area is mostly humpy. The lingual cingulum this one of main cusps. The metaconid is divided to two is developed only. to five cusps, but it predominantly consists of three ones The paraconid of m1s is separated from the metaconid by (mesial metastylid, metaconid s. s., and distal metastylid). a small accessory cusp or a distinct notch. The metaconid The entmetaconid is developed as a crest or a distinct consists of two to three cusps, the last of which is always small cusp. Two cusps form the entoconid, rarely with an the largest. Two various cusps form the entoconid, with accessory cusp either behind the front cusp or behind the an accessory cusp in one case. Between the metaconid back one. The mesostylid is present in four cases only. and entoconid, one or two small accessory cusps are de- The protoconid as a distinct, but blunt cusp is developed veloped. The protoconid is undivided, with an indication at three molars only, rarely with an accessory cusp. The of mesoconid at the back. The hypoconid forms a blunt mesolophid is present in one case and the entprotoconid cusp, with an accessory cusp in front at one molar. Only of three molars protrudes either as a crest or as cusp (-s). two m1s have a cingulum on the lingual side. The blunt hypoconid has rarely developed an accessory 164 Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14, Wien, 2005

Figure 3: Scatter diagram of P4, M1, and M2 measurements of cave bears from the Trojuholník Cave and other ursids from European Pleistocene and Holocene sites (see also tab. 6). cusp in front. The enthypoconid is present at four m2s. developed or indicated on the buccal side only. Tooth decay The posterior crown margin is either without cusps or it is probably present at one m2. consists of some small ones. The cingulum is either not The distinct paraconid of m3s is mostly divided to two SABOL, M., Cave Bears (Ursidae, Mammalia) from … 165

Figure 4: Scatter diagram of m1, m2, and m3 measurements of cave bears from the Trojuholník Cave and other ursids from European Pleistocene and Holocene sites (see also tab. 6). parts, with three accessory cusps at the back in one case. from the both paraconid and entoconid by an accessory Similarly to the paraconid, the metaconid is also mostly cusp (-s). Also entoconid is mostly distinct, rarely undi- developed as a distinct cusp, rarely divided and separated vided cusp with an accessory cusp either in front or at SABOL, M., Cave Bears (Ursidae, Mammalia) from … 166

250 Figure 5: Diagram of the mor- Nixloch 18ka phodynamic indices of P4s and D2 p4s (modified and complemented Lieglloch according to RABEDER, 1992). The question mark indicates the 200 assumed position of the cave bear C2 assemblage from the Trojuholník Nietoperzowa Pod hradem 30ka ? Trojuholník Mixnitz Cave obtained by the analysis of Weinbergh. Erpfingen 150 profile Guloloch l e 66ka 28ka morphodynamic index of p4 .. all P4s (see also tab. 5). oh Vindija G 30ka C1/2 lh 112ka Hohlenstein g e Frauenloch en Schusterlucke Čertova rd Briegl. díra e Zoolith. Spalte Merkenstein the back. The protoconid as H Hartlesgr. 135ka Wildk. Ramesch 64-31ka 35ka a distinct or blunt cusp has 100 Petersh. Drachenl. Sibyllen. mostly developed accessory C1 cusps. The distinct or blunt Hunas 260ka Einhornh. hypoconid is rarely divided Heppenloch to two to three cusps, with the 50 Šipka Jagsthausen enthypoconid in one case. The B1 cingulum is not present.

From all skulls and their frag- morphodynamic index of P4 0 ments, three ones belong to 0 50 100 150 200 250 300 adult individuals, one skull A A B D E belongs to a juvenile, and one fragment is from juvenile to adult individual. Hereby, two skulls probably belong Nine left and seven right branches of mandibles or their to males and one fragment to female. Only two records fragments are present in studied bear fossil material (one represent more or less complete preserved skulls. Other finding consists of the both left and right mandible frag- three are neurocranial parts or their fragments. One skull ments). Four mandibles belong to males and seven prob- (Nr. Z-788) is deformed to a flat shape, causing by the ably to females. Senile individuals are represented by two pressure of sediments. Traces of unknown body impact mandible records, adult individuals by five ones, and one are probably present on other one (Nr. Z-790). mandible belong to juvenile individual. Preserved alveoli

130

ca. 18 ka NL 120

110 27-15 ka Ku2

100 GS 38-25 ka index .. 90 Cu = Conturineshohle .. T? T RK = Ramesch-Knochenhohle RK1 = 0 - 40 cm 80 RK2 = 40 - 60 cm HD3 45-37 ka HD2 HD4-6 Rk3 = 60 - 100 cm RK4 = 100 - 180 cm 70

standardised SO 45-31 ka NL = Nixloch

- 2 Cu 3 SO = Salzofen

p4 .. 60 HD1 44 1 RK 64-31 ka GS = Gamssulzenhohle 4 .. 69 HD = Herdengelhohle 50 87 HD 4-6 = 200-330 cm 110 ka 108 HD 3 = 330-360 cm HD 2 = 360-380 cm HD 1 = 380-420 cm 40 .. Ku2 = Kugelsteinhohle 2 T = Trojuholnik 30 0 10 20 30 40 50 60 70 80 90 100 110 120 P4 - standardised index

Figure 6: Diagram of the morphodynamic standardised indices of P4s and p4s (modified and complemented according to RABEDER, 1995). „T“ with the question mark indicates the assumed position of the cave bear assemblage from the Trojuholník Cave obtained by the analysis of all P4s (see also tab. 5) SABOL, M., Cave Bears (Ursidae, Mammalia) from … 167

of p4s indicate this premolar had only one root in three belong to individuals of cave bears, which are older than cases and in other case, p1 alveolus is probably present fauna of small vertebrates (they come probably from the too. An alveolus of one accessory incisor is situated on Middle Pleniglacial or older period?). They are determined one mandible (Nr. Z-797). to the species Ursus spelaeus ROSENMÜLLER & HEINROTH, In studied sample, eight to nine females have been found, 1794 in spite of the presence of some isolated atavistic whereas other records belong to males, cubs, and individu- characters. als with unknown sex.

5. Acknowledgements 4. Discussion and Conclusion The author is indebted to the Grant Agency for Science, The crown morphology of many studied teeth is typical Slovakia (project No. 1/0080/03) and the Ministry of Edu- for cave bears. However, characters of bears from the both cation, Slovakia (project “Ecosystems of the Late Miocene, Ursus arctos-group (e. g. narrower and shorter anterior Pliocene, and Quaternary – an indicator of the age and cli- parts of M1s, one m1 with a larger back entoconid cusp matic changes”) for financial support. than front one (character observed also at teeth of U. de- ningeri), and four m2s without an accessory cusp between the metaconid and entoconid) and Ursus deningeri-group 6. References (e. g. three m2s with a smaller front entoconid cusp than back one) have also been found at tooth material from the ARGANT, A., 1991. Carnivores quaternaires de Bourgogne. site. Additionally, alveoli of p1s and one-rooted p4s have — Docum. lab géol. Lyon, 115:1–301, Lyon. been preserved at some bear mandibles. BELLA, P. & HOLÚBEK, P., 1999. Zoznam jaskýň na Sloven- From metric point of view, the comparison of measure- sku: 268, Ministerstvo životného prostredia Slovenskej ments of bear teeth from the Trojuholník Cave with republiky, Bratislava. tooth measurements of ursids from the Middle to Late BONIFAY, M.-F., 1971. Carnivores Quaternaires du sud-est Pleistocene European sites (figs. 3 and 4, tab. 6) shows a de la France. — Mém. Mus. Hist. Naturelle, Série C, similarity to Ursus deningeri from Azé I-1 (Argant, 1991), XXI:1–378, Paris. Einhornhöhle (MUSIL, 1972), and Bilzingsleben (MUSIL, COUTURIER, J.A.M., 1954. L’ours brun. — published by the 1991), Ursus spelaeus deningeroides from Azé I-2 and 3 author (Grenoble), 904 p. (ARGANT, 1991), Ursus spelaeus from Saint-Romain (AR- HOLEC, P., OBUCH, J. & POMORSKÝ, F., 1994. Fauna sta- GANT, 1991), Ursus sp. from Okno Cave (SABOL, 2000), vovcov z jaskyne Trojuholník v Borinskom krase v and Ursus priscus from Předmostí (MUSIL, 1964). The Malých Karpatoch. — Slovenský kras, XXXII:71–78, metric similarity of bear teeth from the Trojuholník Cave Liptovský Mikuláš. with records of bears from French sites, especially from HOLEC, P. & POMORSKÝ, F., 1989. Kvartérna fauna niek- Azé I-1, Azé I-2, and Azé I-3, is interesting. torých jaskýň v Malých Karpatoch (Slovensko). From the viewpoint of morphodynamic analysis of fourth — Současný stav a perspektívy výskumu kvartéru v premolars, records of bears from Trojuholník Cave could ČSSR:58–62, Brno. be approximately 30 ka old or even older (figs. 5 and 6). KOBY, E.-F., 1944. Un squelette d’ours brun du pleistocene However, the credibility of that is questionable for reason Italien. — Verh. Naturforsch. Ges., 56:58–85, Basel. of the poor demonstrability of statistic set, when only three MCKENNA, M.C. & BELL, S.K., 1997. Classification of to five P4s and four p4s have been analysed. Also, the exact Mammals: 632 p., Columbia University Press, New circumstances of bear remains findings are unknown. SIL- York. NICKY (1950) only wrote about a find of bear fossils in the MUSIL, R., 1956. Nové osteologické nálezy z jeskyně Axa- depth approximately 10 m during the removal of sediments mitky. — Anthropozoikum, 5:47–54, Praha. from the cave entrance. Besides of them, fossils of micro- MUSIL, R., 1957. Osteologické nálezy zo štěrkovny pod mammals and other small vertebrates (fishes, amphibians, Stránskou skálou, čast 1. — Anthropozoikum, 6:55–69, and birds) have also been found in the cave. However, these Praha. have been found in a thin upper layer (5 to 12 cm) in two MUSIL, R., 1959. Jeskynní medvěd z jeskyně Barové. - of three pits (HOLEC et al., 1994) without fossils of bears or Časopis Moravského musea. — Acta Musei Moraviae, other large mammals (HOLEC & POMORSKÝ, 1989). Fauna XLIV:89–111, Brno. of small vertebrates consists of two assemblages - older MUSIL, R., 1960. Paläontologische Funde in Sedimenten der assemblage of steppe to tundra with Dicrostonyx torquatus, Letzten Zwischeneiszeit. - Časopis Moravského mu- Lemmus lemmus, and Lagopus sp. from the Last Glacial sea. — Acta Musei Moraviae, XLV:99–136, Brno. to the end of the Dryas and younger one of forest-steppe MUSIL, R., 1962. Die Höhle „Švédův stůl“, ein typischer to forest with Clethrionomys sp., Pitymys subterraneus, Höhlenhyänenhorst. — Anthropos, 13(5):97–260, and bats from the start of Holocene (Preboreal to Boreal, Brno. possibly to Atlantic) (HOLEC et al., 1994). MUSIL, R., 1964. Die Braunbären aus dem Ende des letzten On the basis of above mentioned facts, it is possible to Glacials. - Časopis Moravského musea. — Acta Musei draw that fossil records of bears from the Trojuholník Cave Moraviae, XLIX:83–102, Brno. 168 Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14, Wien, 2005

MUSIL, R., 1965. Die Bärenhöhle Pod hradem, Die Entwick- ren aus dem Nixloch bei Losenstein-Ternberg (O.Ö.). lung der Höhlenbären im letzten Glacial. — Anthropos, — Mitt. Komm. Quartärforsch., 8:133–141, Wien. 18(10):7–92, Brno. RABEDER, G., 1995. Evolutionsniveau und Chronologie MUSIL, R., 1972. Die Bären der Stránska skála. — Anthro- der Höhlenbären aus der Gamssulzen-Höhle im Toten pos, 20:107–112, Brno. Gebirge (Oberösterreich). — Mitt. Komm. Quartärfor- MUSIL, R., 1991. Die Bären von Bilzingsleben. — Bilz- sch., 9:69–81, Wien. ingsleben, IV:81–102, Berlin. SABOL, M., 2000. Fosílne a subfosílne medvedŏvité PAULUS, M., 1947. Études sur la faune quaternaire de la mäsožravce (Ursidae, Carnivora) z územia Slovenska: vallée inferieure du Gard ou Gardon. — Bull. Mus. 149 p., Manuscript, Bratislava. Hist. Nat. Marseille, VII, 1:1–30, Marseille. SABOL, M., 2002. Fossil findings of cave bears from the Up- PAUNOVIĆ, M., 1991. Morphometrische und morphogene- per Pleistocene sediments of selected caves in Slova- tische Untersuchungender Zähne des Ursus spelaeus kia. — Mineralia Slovaca, 34(1):35–52, Bratislava. ROSENM. et HEINROTH aus der Höhle Veternica (NW SILNICKÝ, K., 1949. Predbežná správa o náleze zvyškov Kroatien). — RAD Hrvatske akademije znanosti i kostry jaskynneho medvedă (Ursus spelaeus BLMB.) umjetnosti - Razred za prirodne znanosti, 458(25): v pajštúnskom krase (Borinka) v Malých Karpatoch. 1–19, Zagreb. — Príroda, 5(8):120, Martin. RABEDER, G., 1983. Neues vom Höhlenbären: Zur Morpho- SILNICKÝ, K., 1950. Nález zbytkov kostry jaskynného genetik der Backenzähne. — Die Höhle, 34(2):67–85, medvedă (Ursus spelaeus BLMB.) v Pajštúnskom (Bo- Wien. rinka) krase v Malých Karpatoch. — Československý RABEDER, G., 1992. Das Evolutionsniveau des Höhlenbä- kras, 4:211–214, Brno. SABOL, M., Cave Bears (Ursidae, Mammalia) from … 169

TABLES

Table 1: Measurements of upper C sup. C inf. Ursus spelaeus (Trojuholník) and lower canines of cave bears n min. mean max. n min. mean max. from the Trojuholník Cave. longitudinal diameter of crown base 3 20.4 22.9 24.4 4 19.8 24.4 28.8 transverse diameter of crown base 4 14.8 17.2 19.2 4 14.7 18.0 21.4 max. longitudinal diameter of root 2 24.3 28.0 31.6 1 — 23.9 — max. transverse diameter of root 2 15.8 18.2 20.6 1 — 17,8 —

Table 2: Measurements of upper Ursus spelaeus max. length max. width and lower premolars and molars (Trojuholník) n min. mean max. n min. mean max. of cave bears from the Troju- P4 2 19.1 19.3 19.5 3 13.0 13.9 14.5 holník Cave. p4 4 13.2 14.0 14.8 4 9.0 9.3 9.9 M1 4 26.0 28.1 30.8 3 16.5 17.6 18.8 M2 5 40.2 42.6 45.6 4 19.2 20.6 22.9 m1 6 26.3 27.2 29.6 6 13.2 13.8 15.5 m2 10 26.2 28.2 31.2 10 16.4 17.9 19.8 m3 6 22.7 24.8 29.3 5 16.9 18.1 19.3

Table 3: Measurements of cave skulls Ursus spelaeus (Trojuholník) bear skulls from the Trojuholník Z-452 Z-788 Z-790 Z-791 min. - max. mean Cave. max. length 407.0 — — — — 407.0 length of viscerocranial part 309.2 — — — — 309.2 length of neurocranial part 117.1 91.6 110.0 118.4 91.6 - 118.4 109.4 length of palate 220.0 — — — — 220.0 rostral width 98.0 — — — — 98.0 zygomatic width 233.2 — — — — 233.2 interorbital width 85.2 80.0 — — 80.0 - 85.2 82.6 postorbital width 80.4 — 74.0 83.4 74.0 - 83.4 79.3 mastoid width 168.3 — — — — 168.3 height 155.8 — — — — 155.8 forehead width 110.0 91.7 — 138.9 91.7 - 110.0 113.5 max. width of braincase 113.3 — 106.2 113.7 106.2 - 113.7 111.1 length of C sup. - M2 dex. 152.0 — — — — 152.0 length of C sup. - M2 sin. 151.7 128.3 — — 128.3 - 151.7 140.0 length of P4 - M2 dex. 91.6 91.2 — — 91.2 - 91.6 91.4 length of P4 - M2 sin. 90.8 92.8 — — 90.8 - 92.8 91.8

mandibles Ursus spelaeus (Trojuholník) Table 4: Measurements of cave bear mandibles from the Tro- n min. mean max juholník Cave. length 3 274.4 298.2 338.4 max. height 1 — 144.3 — height under m1 8 55.0 60.9 68.7 c inf. - m3 length (alveolar) 6 158.4 170.3 194.7 p4 - m3 length (alveolar) 8 90.5 95.5 106.0 170 Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14, Wien, 2005

a) Ursus spelaeus (Trojuholník) – upper premolars (P4) Table 5a-c: The valuation of morphodynamic Morphotype Amount Factor Product Frequency P4-index P4-stand. indices of cave bear P4s and p4s from the Troju- A/D 1 1 1 33.3 % holník Cave. a – the valuation of unquestionable B 1 1 1 33.3 % P4s, b – the valuation of all P4s, c – the valuation E 1 3 3 33.3 % of all p4s. Σ 3 5 99.9 % 166.67 65.18

b) Ursus spelaeus (Trojuholník) – upper premolars (P4) Morphotype Amount Factor Product Frequency P4-index P4-stand. A? 1 0 0 20 % A/D 1 1 1 20 % A/D? 1 1 1 20 % B 1 1 1 20 % E 1 3 3 20 % Σ 5 6 100 % 120 46.93

c) Ursus spelaeus (Trojuholník) – lower premolars (p4) Morphotype Amount Factor Product Frequency p4-index p4-stand. C1 2 1 2 50 % C3 1 3 3 25 % D1 1 1.5 1.5 25 % Σ 4 6.5 100 % 162.5 81.99

P4 M1 M2 m1 m2 m3 L W L Wf L WP L WH L Wp L WPD Ursus deningeri Einhornhöhle (MUSIL. 1972) 18.8 13.9 26.7 18.7 — — 28.9 14.2 28.8 17.6 25.1 18.4 Urdhöhle (MUSIL. 1972) 17.9 12.4 25.3 17.0 — — 27.0 12.7 27.1 15.9 23.9 14.7 Hundsheim (MUSIL. 1957. 1960. 1972) 18.1 12.8 25.9 18.0 — — — — — — — — Stránska skála (MUSIL. 1972) 18.9 13.7 25.5 18.5 37.9 19.7 27.4 12.8 26.9 16.8 23.0 17.2 Bilzingsleben (MUSIL. 1991) 18.4 13.2 26.1 18.2 42.4 21.3 28.0 13.9 29.1 17.7 26.0 18.4 Chateau (ARGANT. 1991) 18.2 13.2 — — 42.9 21.3 — — 29.2 17.8 26.1 18.5 Azé I—1 (ARGANT. 1991) 18.7 13.2 — — 43.1 21.4 — — 28.5 17.7 24.2 17.9 Ursus spelaeus deningeroides Azé I—2 (ARGANT. 1991) 19.4 14.2 — — 44.1 23.1 — — 27.9 17.4 25.0 17.9 Azé I—3 (ARGANT. 1991) 18.8 13.5 — — 40.9 21.3 — — 28.0 17.3 24.3 18.0 Ursus spelaeus Berze (ARGANT. 1991) 17.8 13.2 — — 40.9 21.5 — — 28.7 17.6 26.1 18.5 Saint—Romain (ARGANT. 1991) 19.1 13.8 — — 43.2 21.6 — — 28.8 17.1 25.7 18.1 Saint—Aubin (ARGANT. 1991) 18.4 13.2 — — 42.5 20.9 — — 29.2 16.7 25.6 18.0 Grottes du Gardon (PAULUS. 1947) 20.0 14.8 — — 45.3 23.3 — — — — 27.0 18.4 Veternica (PAUNOVIĆ. 1991) 20.3 14.0 — — 44.9 23.2 — — — — 27.5 19.7 Švédův stůl (Eemian—Brőrup) (MUSIL. 20.6 14.0 29.2 19.7 46.1 23.1 31.1 14.0 32.0 18.9 28.1 19.7 1962) Švédův stůl («W2») (MUSIL. 1962) 20.9 14.5 29.1 19.5 46.4 22.5 31.1 14.8 31.1 19.0 28.0 20.4 Pod hradem I (MUSIL. 1965) 19.8 14.5 29.2 20.3 44.4 22.4 30.0 14.2 31.0 18.3 26.7 18.9 Pod hradem II (MUSIL. 1965) 20.8 14.4 29.0 19.6 45.7 23.3 30.7 14.6 31.5 18.5 28.1 19.8 Pod hradem III (MUSIL. 1965) 20.7 14.8 27.8 18.6 44.8 22.5 30.7 14.9 31.2 18.5 27.7 20.0 Pod hradem IV (MUSIL. 1965) — — — — — — 29.4 14.2 29.9 18.4 27.2 19.1 Barová (MUSIL. 1959) 19.4 13.4 28.5 19.7 44.5 22.7 29.8 14.7 30.0 18.4 27.6 19.5 Aksamitka (MUSIL. 1956) — — — — — — — — — — 28.0 20.0 Malá Ladnica — — — — — — 27.2 13.8 27.2 18.1 24.3 17.1 Bezmenná jaskyňa v Malom vrchu — — — — — — — — 30.0 19.0 — — Činčianska jaskyňa 19.0 13.6 27.3 18.3 45.4 20.7 28.9 14.2 29.5 18.0 26.6 18.9 Domica — — 27.4 18.4 44.2 24.5 28.4 14.6 28.9 19.2 27.7 18.2 SABOL, M., Cave Bears (Ursidae, Mammalia) from … 171

P4 M1 M2 m1 m2 m3 L W L Wf L WP L WH L Wp L WPD Horná Túfna 19.9 15.3 — — 44.8 22.6 30.6 14.4 30.5 18.3 27.5 18.3 Jánošíkova jaskyňa v Šulovských skalách 21.9 15.2 31.3 21.7 46.9 24.7 31.4 14.7 32.8 19.2 28.2 20.4 Jasovská jaskyňa 20.5 13.9 28.1 19.8 47.3 22.6 31.2 15.4 — — 25.2 17.6 Malá drienčanska jaskyňa — — — — — — 33.2 16.0 27.6 19.2 — — Medvedia jaskyňa pod Sivým vrchom 18.6 13.5 27.1 17.4 42.6 20.4 32.4 15.7 31.7 20.4 — — Ursus spelaeus Medvedia jaskyňa v Slovenskom raji 20.6 14.6 28.7 19.8 43.1 22.9 30.5 13.5 30.8 18.6 27.9 19.6 Moldavská jaskyňa — — — — — — — — 32.8 19.8 24.0 20.7 Psie diery 21.5 15.5 32.1 21.2 46.4 24.8 28.7 14.0 29.6 18.5 28.4 20.4 Tmavá skála 20.2 13.7 30.3 19.0 45.2 21.9 30.6 14.5 31.3 18.3 26.7 18.9 Važecka jaskyňa — — 28.3 19.6 45.2 22.4 30.2 14.6 30.4 18.6 26.8 18.7 Trojuholník 19.3 13.9 28.1 17.3 42.6 20.6 27.2 13.8 28.2 17.5 24.8 18.1 Ursus ingressus Gamssulzenhöhle 1 (RABEDER. 1995) 20.1 14.2 28.7 19.8 44.4 22.6 — — 30.6 18.3 27.6 19.1 Gamssulzenhöhle 2 (RABEDER. 1995) 19.6 13.7 28.5 19.5 43.5 22.7 — — 29.5 18.0 26.6 18.9 Gamssulzenhöhle 3 (RABEDER. 1995) 20.3 14.6 28.3 20.0 43.9 22.7 — — 30.0 18.8 27.7 19.3 Ursus sp. Okno 19.4 13.9 26.7 17.6 44.1 20.9 28.5 13.5 29.5 17.6 26.9 18.0 Ursus prearctos Grimaldi (BONIFAY. 1971) — — — — — — — — — — 19.1 14.1 Ursus (cf.) priscus Maspino (KOBY. 1944) 17.0 13.4 — — 39.2 19.1 — — — — — — Předmostí (MUSIL. 1964) 15.0 9.0 24.2 17.4 38.3 20.6 27.1 14.2 28.8 17.9 24.2 17.3 Važecká jaskyňa 17.2 12.6 21.6 16.0 33.7 18.4 — — — — — — Vyvieranie 22.4 13.8 31.7 21.7 46.5 21.6 — — — — — — Kupčovie izbička 16.3 12.4 25.2 16.4 38.3 19.0 25.7 12.4 25.5 16.8 Liskovská jaskyňa 17.6 13.3 24.0 15.9 38.4 18.0 — — — — — — Ursus arctos Flavigny (ARGANT. 1991) 18.0 13.6 — — 38.4 20.3 — — — — 22.5 16.7 Moldavská jaskyňa — — — — — — 26.8 13.2 28.0 17.2 23.3 16.6 Ursus arctos arctos Pyrenees (COUTURIER. 1954) — — — — — — — — 22.0 13.5 17.5 13.5 Líščia jaskyňa — — — — — — — — 26.8 16.1 — — Klenovec — — — — — — 23.4 12.6 24.2 15.6 — — Lukáčova priepast 17.3 13.0 23.3 14.6 36.8 18.9 — — 26.1 17.6 19.4 15.4 Medvedia jaskyňa — Hučiaky 13.8 11.3 19.9 12.5 29.0 15.6 21.5 10.0 23.2 13.4 17.3 12.8 Šípová jaskyňa 15.4 10.1 21.2 14.1 — — — — — — — — Očová 15.1 10.4 20.7 15.0 33.2 16.2 — — 22.7 14.3 19.4 14.8

Table 6: The premolar and molar measurements of bears from European Pleistocene and Holocene sites (see also figs. 3 and 4). L – tooth length, W – tooth width, Wf – width of the front part, WH – width in the place of hypoconid, WP – width in the place of paracone, Wp – width of the posterior part, WPD – width in the place of paraconid. 172 Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14, Wien, 2005

TABLE 1

Ursus spelaeus ROSENMÜLLER & HEINROTH, 1794; skulls, Trojuholník Cave, photo: L. Osvald

Figs. 1-3: Z-452

Figs. 4-6: Z-788 (deformed skull); 1 and 5 – dorsal view, 2 and 6 – ventral view, 3 and 4 lateral view. SABOL, M., Cave Bears (Ursidae, Mammalia) from … 173

TABLE 1 174 Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 14, Wien, 2005

TABLE 2

Ursus spelaeus ROSENMÜLLER & HEINROTH, 1794; mandibles, Trojuholník Cave, photo: L. Osvald

Fig. 1: right mandible with m2 (Z-798).

Fig. 2: right mandible of a juvenile with m3 (Z-1001) in crypt.

Figs. 3-4: left mandible with m1-2 (Z-797).

Figs. 5-6: right mandible with m1-3 (Z-800); 1, 4 and 6 – buccal view, 2 – lingual view, 3 and 5 – occlusal view. SABOL, M., Cave Bears (Ursidae, Mammalia) from … 175

TABLE 2