- Email: [email protected]
Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians
Quaternary International xxx (2013) 1e8
Contents lists available at SciVerse ScienceDirect
Quaternary International journal homepage: www.elsevier.com/locate/quaint
Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians Martin Sabol a, *, Andrej Bendík b, Michal Grivalský c, Martin Grivalský c, Ján Lizák c, Igor Michlík c a b c
Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, SK-84215 Bratislava, Slovakia Slovak National Museum in Martin e Museum of Andrej Kmet, A. Kmet Street 20, SK-03601 Martin, Slovakia Spisská Belá Speleological Group, kpt. Nálepku Street 23, SK-059 01 Spisská Belá, Slovakia
a r t i c l e i n f o
a b s t r a c t New research of the Izabela Textorisová Cave in the Velká Fatra Mts. and the Javorinka Cave in the Vysoké Tatry Mts. yielded important data on Western Carpathians cave bear palaeopopulations. The radiometric dating of cave bear fossils from the Javorinka Cave (c. 51,000 14C BP) represents one of the oldest documented record of cave bears in Slovakia. The site is also the highest documented occurrence of cave bears with an altitude more than 1500 m. The radiocarbon dates from the Izabela Textorisová Cave (30,181 e30,190 cal. BP) represent the last known occurrence of cave bears in the Slovakian Western Carpathians. Ó 2013 Elsevier Ltd and INQUA. All rights reserved.
Article history: Available online xxx
1. Introduction The fossil remains of cave bears (Ursus ex gr. spelaeus) from the Slovak territory are known at least since the Middle Ages. The first mentions of these “dragon bones”, however, occur in historical documents of the seventeenth and eighteenth century (Hain, 1672; Marsigli, 1726; Bell, 1736; Brückmann, 1739). The English traveller Townson (1797) was the first who distinguished the bear fossils among “bones of dragons” from the Demänová Caves. The beginning of more scientific research of cave bear fossil record from Slovak caves can be dated from that time. The first detailed morphological and metrical descriptions of cave bear fossils from the territory under study were published after World War II (Musil, 1953, 1956; Janácik and Schmidt, 1965). To date, fossil remains of cave bears are known from at least 60 Slovak caves (Sabol, 2001), although this number could be higher than 100 sites. With international scientific cooperation with Vienna University and the Austrian Academy of Science, a modern systematic research of some important sites started in 2007 (Sabol et al., 2008b; Laughlan et al., 2012), accompanied also by AMS dating of cave bear fossils (Ábelová and Sabol, 2009). Simultaneously, exploration of other regions expected to have a new cave bear fossil record have been undertaken in highland and alpine
* Corresponding author. E-mail addresses: [email protected] (A. Bendík).
(M.
Sabol),
[email protected]
areas of the Velká Fatra Mts. (Bendík, 2005) and the Vysoké Tatry Mts. (Plu cinský, 2011). Subsequent preliminary study and dating of osteological remains from two sites (Izabela Textorisová Cave in the Velká Fatra Mts. and Javorinka Cave in the Vysoké Tatry Mts.) has produced interesting data on the last occurrence and the geographical distribution of cave bears in the Slovakian Western Carpathians. 2. Localities The Izabela Textorisová Cave is situated in the Konský dol Vale within the Gaderská Valley in the Tlstá National Natural Reservation, Velká Fatra Mts., northern Slovakia (Fig. 1). The site is formed by the Gader limestone (dark-grey compact limestone and lightgrey crinoid limestone, Upper Anisian, Middle Triassic) of the Choc Nappe (Hronikum Unit). The cave consists of a large portal entrance and three halls, more or less connected by horizontal corridors. The bottom of the halls is mostly covered by coarse debris with loam or with layers of guano (thickness to 20 cm). The main location of the discovery of cave bear fossils (named “Hrobka”, Tomb) is situated at the end of one tectonically steep inclined ” (Hall). This corridor is closed corridor in the cave section of “Sien by a cave-in (or collapse), which probably obstructed the original access path for bears. The sedimentary sequence in cave section “Hrobka” consists of an overlying gravel accumulation (covered by 2 cm thick layer of white flowstone), a loamy layer with larger clasts (from 2 cm to 8 cm) and scattered rocks (up to 40 cm in size), and with a thick layer of light-grey clay underlying. The bear fossils have
1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.05.022
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
2
M. Sabol et al. / Quaternary International xxx (2013) 1e8
Fig. 1. Location of the Izabela Textorisová Cave in the Velká Fatra Mts., northern Slovakia.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
M. Sabol et al. / Quaternary International xxx (2013) 1e8
3
at an altitude of 1559 m, with height 7e8 m and width up to 5 m, oriented parallel to the Kolová Valley (Plu cinský, 2011).
been found mainly at depths from 30 to 80 cm above a wet clayey layer. At the same depth, more or less complete skeletons of four to nine cave bear specimens have also been discovered in situ position (Sabol et al., 2008a). ”, a Cave bear remains have also been found in the “Sien 200 300 150 cm section situated at the right side behind the cave entrance. Its base consists of up to 5 cm large clasts and rare large rock blocks mixed with unstratified layers of cave loam, partly with a speleothem. Occasionally, faintly rounded pebbles also occur here, indicating a sporadic underground river. A cave bear record from this cave section (to date, more than 500 fossil remains) consists mainly of teeth, phalanges, and vertebrae. The limb bones are represented in smaller quantities, together with rare skulls and mandible fragments. Some bones show gnawing marks (caused by hyenas?), or fractures (caused by humans?). Javorinka Cave (Fig. 2) is situated in the western part of the Kolový Úplaz Massif (1783 m) on a western slope of the central Javorová Valley on the northern side of the Vysoké Tatry Mts., close to 1254 m elevation (Fudaly et al., 1983; Plu cinský, 1996). The entrance of this corrosive-erosional cave, discovered in 1973 (Fudaly et al., 1983), is located at an altitude of 1218 m (Plu cinský, 1996). The cave is a part of the JavorinkaeNádejná cave system with a current length of 10,501 m (Plucinský, 2011) and consists of carbonate rocks of the Tatrikum Unit (Lower Triassic dark-grey to dark-red sandy laminated clayed shale, Middle Triassic grey dolomite and Guttenstein limestone, light-grey dolomite and rose dolomite with intercalations of hematite, Middle Jurassic red crinoid limestone and grey-rose to grey clayey limestone, and Malm-Neocomian limestone) with tectonic faults oriented NWeSE and NEeSW (Fudaly et al., 1983). Most horizontal and inclined cave
3. Fossil material Fossils of cave bears from the Izabela Textorisová Cave were discovered in 2004 and excavated gradually up to 2007 (Bendík, 2005, 2007; Bendík and Sabol, 2007). More than 1500 bear remains have been found, and are housed in the Slovak National Museum e Museum of Andrej Kmet in Martin town. The fossil record from the site is now under study, and only data on some pathological phenomena (Sabol et al., 2008a; Stuller et al., 2011) with a first attempt of AMS dating (Bendík and Sabol, 2007) are published. To obtain more data on the radiocarbon age of fossils from the cave, some metapodial bones from the “Hrobka” and the ” cave parts were submitted to the VERA Laboratory of Vienna “Sien University. Two of these samples (Mc IV dext. and Mt V sin., described below) yielded numerical dates for the latest occurrence of cave bears in the Slovak territory of the Western Carpathians. Mc IV dext. (HJ/1156) from “Hrobka” cave part is a juvenile or subadult(?) specimen. The grey-white bone without the unjoined distal epiphysis was damaged faintly on its surface and shows no pathological phenomena or marks of post-mortem agents. The ” was adult yellowebrown to brown Mt V sin. (HJ/S-209) from “Sien also damaged on the surface and without pathological phenomena or marks of post-mortem agents (Fig. 3e,f). The measurements of both metapodials are given in Table 1. The fossil record of cave bears from the site is preliminarily assigned to Ursus ex gr. spelaeus. Isolated remains of wolf (Canis lupus) and cervids (cf. Capreolus capreolus, Cervus sp.) have also been found.
Table 1 Measurements (in mm) of cave bear metapodial bones from both studied Slovak caves. Metacarpalia
Maximum length Max. transversal diameter of the proximal epiphysis Max. dorso-plantar diameter of the proximal epiphysis Minimum transversal diameter of the diaphysis Minimum dorso-plantar diameter of the diaphysis Max. transversal diameter of the distal epiphysis Max. dorso-plantar diameter of the distal epiphysis Width of the distal head Metatarsalia
Maximum length Max. transversal diameter of the proximal epiphysis Max. dorso-plantar diameter of the proximal epiphysis Minimum transversal diameter of the diaphysis Minimum dorso-plantar diameter of the diaphysis Max. transversal diameter of the distal epiphysis Max. dorso-plantar diameter of the distal epiphysis Width of the distal head
Izabela Textorisová C. e Hrobka
Javorinka Cave
Mc IV dext. (HJ/1156)
Mc V dext. (HJ/1485)
Mc I dext.
Mc II dext.
Mc II sin.
Mc III sin.
Mc V dext.
e 21.3 31.0 16.4 14.2 e e e
75.5 26.0 28.2 16.2 13.65 23.4 e 15.9
57.5 22.05 16.39 12.14 9.91 19.15 17.0 18.01
70.93 19.09 24.97 17.42 12.41 25.07 18.71 19.55
e 16.52 22.68 14.63 11.1 e e e
80.76 22.54 32.05 16.63 15.21 26.83 22.94 21.91
78.55 28.41 30.99 18.99 16.99 28.19 19.19 25.98
Izabela Textorisová Cave e Hrobka
Izabela Textorisová e Sien
Javorinka Cave
Mt IV sin.
Mt V sin.
Mt III sin.
Mt IV dext.
Mt V sin.
Mt III sin.
(HJ/1345)
(HJ/1343)
(HJ/S-361)
(HJ/S-208)
(HJ/S-209)
79.1 20.2 30.1 17.6 13.2 19.6 e 14.6
92.4 22.8 35.5 18.0 17.5 27.0 e 21.9
83.0 e 31.2 16.5 14.5 24.9 e 19.6
e 19.4 18.4 14.1 12.2 e e e
e 23.4 26.9 11.2 13.0 e e e
corridors formed by river activity, and these are characterized by the abundant occurrence of granitoids, especially in the lower ones. This granitoid material represents alluvial deposits of Kolový Creek, deposited during the Pleistocene gradual incision of the Kolová Valley. In older corridors, the gravel deposits are covered by sandy loam, limestone debris, or thin flowstone. Cave bear fossils have been found in two cave parts: “Medvedí dóm” (Bear Dome) and the “Medvedia chodba” (Bear Corridor). Whereas the “Medvedí dóm” represents a relatively large, 30 m high cave space at an altitude of 1525 m, the “Medvedia chodba” is a 150 m long horizontal corridor
(HJ/1344) 83.2 28.4 29.5 14.2 15.6 25.7 18.2 22.4
84.04 21.79 36.59 16.52 12.65 22.89 e 20.28
Mt IV sin.
Mt V sin.
81.61 21.64 29.35 16.25 13.22 23.19 17.6 20.36
83.66 27.48 30.28 12.74 13.47 24.69 16.86 21.59
Cave bear fossils from the Javorinka Cave were discovered during speleological exploration in 2011 (in the “Medvedí dóm”) and 2012 (in the “Medvedia chodba”) directly on the surface of the cave base (Fig. 4). During sampling of cave bear fossils in the “Medvedí dóm” in summer 2011, a fragment of right maxilla with dentition (C e M2 dext.), I3 dext., p4 dext., m1 dext., and m2 sin. together with metapodials (Mc IeII and Mc V dext., Mc IIeIII sin., Mt IIIeV sin. (of one specimen), and two fragments) and four phalanges were collected. Two groups of bones (metapodials and phalanges) were sent to Vienna VERA-Laboratory for AMS dating.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
4
M. Sabol et al. / Quaternary International xxx (2013) 1e8
The maxilla fragment without P3 (Fig. 3a) is partly covered by a filmy speleothem layer and faintly damaged. The light crowns of preserved cheek teeth are unworn (Stiner’s wear stage III; Stiner, 1998), indicating a young specimen, probably male on the basis of the canine size. The P4 crown morphology resembles Rabeder’s (1999) morphotype F with tiny accessory morphological elements on inner sides of both the metacone and the protocone as well as between the protocone and the paracone. The crown morphology of M1 with the cingulum terminated under the hypocone fully corresponds with Rabeder’s morphotype C3. The M2 crown, with the cusp-shaped cingulum extended as far as the posthypocone, also shows an advanced stage of morphology in the parastyle complex (C3), the mesostyle complex (C), the talon (C/D) as well as in the paracone and protocone inner side morphology (C and 2). Crowns of isolated teeth are also unworn (juvenile specimens) and broken roots of these are partly rounded by water. The morphology of p4 crown (Fig. 3b) indicates Rabeder’s morphotype E with the distinct metalophid, but without conspicuous hypoconid and entoconid. The m1 crown (Fig. 3c) with two mesial metastylids, 4-cusped entoconid complex (B4), 2-cusped enthypoconid, and the distinct hypoconulid separated from the entoconid by one tiny
Brownish metapodial bones of adults (Table 1) are mostly undamaged, and show no pathologies and traces of post-mortem agents (apart of Mc II sin. with probably bitten-off distal epiphysis). Apart from cave bear remains (preliminarily attributed to Ursus ex gr. spelaeus), fossils of Gulo gulo and Martes martes have also been found in the “Medvedí dóm”. 4. Results Uncalibrated radiocarbon dates are given as 14C BP and calendar dates as cal. BP (calculated with OxCal3.10 and the INTCAL09 calibration curve). The first radiocarbon dating of a cave bear left radius from the Izabela Textorisová Cave yielded >45,000 14C BP (Bendík and Sabol, 2007). For that reason, some new samples of cave bear bones were dated by AMS in the VERA Laboratory in Vienna (Table 3). Surprisingly, two new dates demonstrate that the Izabela Textorisová Cave was inhabited by cave bears before c. 24,000 14C BP (c. 30,000 cal. BP). This indicates one of the latest occurrence of cave bears in the Slovakian territory of the Western Carpathians. Thus, the last palaeopopulations became extinct here before the beginning of the Last Glacial Maximum (LGM, c. 30,000 years ago).
Table 3 Radiometric dates of cave bear bone samples from both studied Slovak caves by 14C-AMS in the VERA-Laboratory, Vienna (Austria). Data were calculated with OxCal3.10 and the INTCAL09 calibration curve and correspond to the 2s-confidence level. Probability of the individual time periods is in brackets. For calibration, the pMC (percent Modern Carbon) input option was used (corresponding to a symmetric uncertainty of the uncalibrated 14C age). Site
Lab.-Nr.
Sample name
d13Ca [&]
Uncal. BP age range (2s)
cal. BP age range (2s)
Izabela Textorisová Cave Izabela Textorisová Cave Izabela Textorisová Cave Javorinka Cave
VERA-4041 VERA-5679 VERA-5680 VERA-5682
Radius sin. Mc IV dext. (HJ-1156) Mt V sin. (HJ/S-209) Metapodials
19.5 23.4 23.0 20.7
>45,000 24,640 170 26,460 180 51,000 þ 4500/2900
30,181 cal. BP (95.4%) 29,020 cal. BP 30,190 cal. BP (95.4%) 28,970 cal. BP
a
1.2 0.6 5.5 0.9
1s-error.
accessory cusp represents more advanced morphological stage. Morphology of the m2 crown (Fig. 3d) is relatively advanced, showing a higher level of evolution in the metalophid and metastylid complex, bifurcated mesolophid, 4-cusped entoconid complex, or in the enthypoconid complex (C/D). The measurements of all teeth are given in Table 2.
Table 2 Measurements (in mm) of cave bear teeth from Javorinka Cave in the Vysoké Tatry Mts. L e maximum crown length (antero-posterior dimension), TAW e talonid width, TRW e trigonid width, W e maximum crown width (transverse dimension), WA e anterior crown width, WP e posterior crown width. I3 dext.
L W
15.06 14.87
C1 dext.
L W
25.85 21.49
P4 dext.
L W
21.54 15.28
M1 dext.
L WA WP
29.59 20.14 20.76
M2 dext.
L WA WP
45.90 24.94 23.14
p4 dext.
L W
15.47 10.42
m1 dext.
L TRW TAW
30.15 11.77 14.03
m2 sin.
L TRW TAW
32.91 19.24 19.98
Dating of cave bear samples (metapodials and phalanges) from Javorinka Cave is close to the limit of 14C-dating (Table 3). The date indicates inhabitation of the site by cave bears at least in the Middle Weichselian before 50,000 14C BP. Fossil remains of cave bears found between 1525 and 1559 m in the Javorinka Cave, however, represent the highest occurrence in Slovakia, and can be regarded as a record of a high alpine cave bear palaeopopulation from early MIS 3.
5. Discussion Cave bears from the spelaeus-group (including at least two lineages e Ursus spelaeus sensu scripto and Ursus ingressus) evolved in Europe more than 100,000 years ago (Curry, 2010). The transition between these and their precursors from the deningeri-group is generally considered to have occurred at around the beginning of the Last Interglacial (Rabeder et al., 2000). They were adapted to a wide range of environments, but also occupied different ecological niches (Knapp et al., 2009), in an extreme continental climate. Their established distribution extends eastwards from northwest Spain (Grandal d’Anglade, 1993) across Central Europe and the Urals (Kosinstev, 2007) up to the Altai (Knapp et al., 2009), and from Belgium and the Hartz region of Germany in the north (Rabeder et al., 2000) as far south as Italy (Minieri et al., 1995) and Greece (Kostopoulos, 2006), and southeast to the Crimea (Nordmann, 1858e1860; Encloe et al., 2000). The most northerly record of cave bears is from the northern Urals (62 N; Knapp et al., 2009) and the highest one is from the Italian Conturines Cave, with its entrance at 2800 m above sea level (Rabeder et al., 2006). This high alpine occurrence could be possible only during a warmer Last Glacial period, as the expansion of glaciers during the LGM would have made most mountain areas uninhabitable (Pacher and Stuart, 2009).
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
5
Fig. 2. Javorinka Cave in the Vysoké Tatry Mts., northern Slovakia. Locations where cave bear fossils have been found are marked by ellipses.
M. Sabol et al. / Quaternary International xxx (2013) 1e8
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
6
M. Sabol et al. / Quaternary International xxx (2013) 1e8
Fig. 3. Analysed cave bear fossils from Javorinka Cave (aed) and Izabela Textorisová Cave (eef). a e Right maxilla fragment with C and P4-M2, b e p4 dext., c e m1 dext., d e m2 sin., e fragment of metacarpal bone Mc IV dext. (HJ/1156), f e metatarsal bone Mt V sin. (HJ/S-209); metapodial bones were used for radiocarbon dating.
Cave bears became extinct in northern Eurasia and elsewhere during the last GlacialeInterglacial cycle (Pacher and Stuart, 2009), approximately 25,000 years ago (Curry, 2010). According to former assumptions (Kurtén, 1958, 1976; Musil, 1981), the cave bear could survive in some suitable areas well beyond the LGM, i.e. into the Late Glacial and Holocene, implying their survival until at least c.
12,000e15,000 14C BP. Pacher and Stuart (2009), however, found no evidence that cave bears survived into the Late Glacial in any region. The uncalibrated age data of cave bear remains younger than 30,000 14C BP are known mainly from Balme à Collomb (France), Bärenloch (Switzerland), Lieglloch, Gamssulzen, Ochsenhalt (Austria), and Potocka zijalka (Slovenia). Based on the calibrated data, only the records from Balme à Collomb and Gamssulzen are younger than 30,000 cal. BP. (Fiebig and Pacher, 2006). Cave bears had disappeared in Alps and adjacent areas by about 24,000 14C BP (c. 27,800 cal. BP) (Pacher and Stuart, 2009). Within Central Europe, this datum coincides fairly closely with the cooling at the onset of Greenland Stadial 3 (GS-3) (c. 27,500 cal. BP) (Lowe et al., 2008), well before the LGM, most likely attributable to a marked deterioration in quantity and quality of available plant food (Pacher and Stuart, 2009). Cave bear extinction could thus be plausibly linked to the cooling climate and vegetational changes in GS-3 and GS-2 (broadly “LGM”) (Pacher and Stuart, 2009). On the other hand, climate changes cannot be solely to blame for this event, although the prolonged cold period (LGM) shortened and eliminated growing seasons and changed plant distributions across Europe (Curry, 2010). Though there is evidence of cave bear hunting by ancient hominids (Neanderthals and/or modern humans), it is not likely to have led to the extinction of these animals (Curry, 2010). Rather, different hibernation strategies of cave and brown bears might made a difference (Stiller et al., 2010). Judging from the relative amounts of cave bear and brown bear remains in European caves, cave bears were more dependent on caves for hibernation (Kurtén, 1976; Rabeder et al., 2000). Both modern humans and Neanderthals would have been strong competitors for these caves, and might have forced cave bears into less suitable sites for hibernation (Grayson and Delpech, 2003), where cave bear cubs were more in danger of killing by predators and/or cave bear males. According to the latest aDNA study (Knapp et al., 2009; Stiller et al., 2010), cave bear palaeopopulations began a long, slow decline 50,000 years ago (i.e. approximately 25,000 years before their extinction), around the Middle-to-Upper Palaeolithic transition and well before the LGM began (Curry, 2010). This slow demise
Fig. 4. Fossil accumulations in the Medvedí dóm part of Javorinka Cave, whence some specimens have been collected for analyses.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
M. Sabol et al. / Quaternary International xxx (2013) 1e8
of cave bears suggests that one or more environmental factors must have subtly increased their mortality and/or decreased their reproduction rate (Stiller et al., 2010), although the range of their geographical distribution was not changed (Musil, 1981, 1986). According to Knapp et al. (2009), the cave bear evolutionary history before their extinction involved a rather complex phylogeographic pattern and a more substantial loss of biological diversity at the end of the Pleistocene than has been previously recognized. In any case, their disappearance corresponds broadly with the withdrawal of many extinct and extant mammal species from large area of Europe around the LGM (Pacher and Stuart, 2009). Within the Slovakian territory of the Western Carpathians, the vertical distribution range of cave bears was from 215 m a.s.l. (the Cave) (Sabol, 2001). The Cin cany Cave) to 1559 m a.s.l. (the Murán fossil record from the latter site, however, is documented only by a find of weathered ursid canine (Fejfar, 1973), with attribution to a cave bear highly questioning. Thus, the fossils of cave bears in the “Medvedí dóm” in the Javorinka Cave represent their highest recorded occurrence in the Slovakian Western Carpathians, 1525e 1559 m a.s.l. Although no systematic research of the site has been realized to date and bear remains were preliminary determined only as Ursus ex gr. spelaeus, the advanced morphology of the molars together with a comparison of the M2’s length in relation to the cave part altitude (cf. Horacek et al., 2012: Fig. 2) indicate attribution to the taxon U. ingressus, the presence of which has been validated in Slovakia during a research of Medvedia Cave in the Slovenský raj Mts. (Sabol et al., 2008b). This high alpine occurrence
7
of bears was only enabled by favourable climatic conditions during a relatively warmer Last Glacial period at the beginning of MIS 3 (Ábelová and Sabol, 2009). The date 51,000 þ 4500/2900 14C BP represents one of the oldest documented record of cave bears in the territory under study. The range of the uncalibrated radiocarbon dates from Slovak cave bear fossils extends from 39,100 þ 1100/1000 14C BP (Ceklovský and Sabol, 2012) to 51,100 þ 4700/2900 14C BP (Sabol et al., 2008b), with unaccepted dates of >15,000 14C BP from Medvedia Cave in the Slovenský raj Mts. (Schmidt and Chrapan, 1970) and 15,490 780 to 17,530 900 14C BP from Psie diery Cave (Pomorský, 1993). New radiometric dates from Izabela Textorisová Cave, however, move the upper level of this range to 24,640 170 14C BP or 26,460 180 14C BP (30,181 cal. BP. and 30,190 cal. BP), approximately coincident with a period between Greenland Stadial 4 and 5. These can be assumed as approximate dates of the last occurrence of cave bears in the Slovakian Western Carpathians, with extinction before the LGM. This is in relatively good agreement with results from the Alps and adjacent areas, where cave bears had disappeared by about 27,000e28,000 cal. BP. Fossils of cave bears are entirely absent in Slovakian Upper Palaeolithic sites (e.g. Vla ciky, 2009), whereas sediments of Neanderthal Middle Palaeolithic sites contain these, together with remains of Ursus arctos. Preliminary results from carbon isotopes (d13C food from 28.6& to 22.6&) indicate that the bears fed on plants in a steppe environment, alternating with meadows and light or dense
Fig. 5. Relations among uncalibrated radiocarbon ages of cave bear fossils from Slovak cave sites, altitude of these caves, and isotopic data (d13C). Although there are so far only a few numerical data, a trend of the cave bear palaeopopulations shift to lower (more forested) altitudes with coming LGM can be assumed (data source: Bendík and Sabol, 2007; Sabol et al., 2008b; Ábelová and Sabol, 2009; Ceklovský and Sabol, 2012; Laughlan et al., 2012; and unpublished data).
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
8
M. Sabol et al. / Quaternary International xxx (2013) 1e8
taiga-like forests (Ábelová and Sabol, 2009). Based on available data, there is also a possibility to suggest movement of cave bear palaeopopulations from high alpine areas to low-altitude (more forested) habitats during the gradually climatic cooling of the coming LGM (Fig. 5). 6. Conclusions The new results of radiometric dating of cave bear fossils from two Slovakian caves (Izabela Textorisová Cave in the Velká Fatra Mts. and Javorinka Cave in the Vysoké Tatry Mts.) yielded important data on Western Carpathians palaeopopulations of this ursid. The date from Javorinka Cave (51,000 þ 4500/2900 14C BP) represents one of the oldest documented record of cave bears in the territory of Slovakia, and can be correlated with the beginning of MIS 3. The site also represents the highest fossil record of cave bears for the area under study with an altitude between 1525 and 1559 m. The new radiocarbon dates from Izabela Textorisová Cave (30,181 cal. BP. and 30,190 cal. BP) represent the last known occurrence of cave bears in the Slovakian Western Carpathians. Palaeopopulations became extinct before the LGM began, what is in good agreement with data from other Central European regions. Acknowledgements The research for this paper was carried out with financial support from the Slovak Research and Development Agency under contact APVV-0280-07 and the Slovak Grant Agency under contract Vega 1/0396/12. The authors also gratefully acknowledge the helpful comments and suggestions of Dr. Evelyne Crégut-Bonnoure and Dr. Silviu Constantin. Their gratitude goes also to Dr. Norm R. Catto for English language review. References Ábelová, M., Sabol, M., 2009. Attempt on the reconstruction of palaeotemperature and palaeoenvironment in the territory of Slovakia during the Last Glacial based on oxygen and carbon isotopes from tooth enamel and bone collagen of cave ovská, Z., Vl bears. In: Sabol, M., Visn cek, L. (Eds.), European Cave Bear Researches. Slovenský kras 47 (Suppl. 1), 5e18. Liptovský Mikulás. Bell, M., 1736. Notitia Hungariae novae historico geographica, divisa in partes qvatvor, quarum prima, Hungariam Cis-Danubiam; altera, Trans-Danubiam; tertia, Cis-Tibiscanam; quarta, Trans-Tibiscanam. Tomus Secundus, Vienna Austriae. p. 579. jaskynného (Ursus spelaeus Bendík, A., 2005. Nové osteologické nálezy medveda Rosenmüller et Heinroth) vo Velkej Fatre. Zborník Kmetianum X, 226e231. Bendík, A., 2007. Zachovanie kostier medveda jaskynneho (Ursus spelaeus Rosenmüller, 1794) v jaskyni Izabely Textorisovej vo Velkej Fatre. Slovensky kras XLV, 219e230. Bendík, A., Sabol, M., 2007. Cave bears from the Cave of Izabela Textorisová (the Velká Fatra Mts., Slovakia) e a state of the art. Scripta Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis 35, 151e156. Geology. Brückmann, F., 1739. Antra draconum Liptovensis. Epist. itineraria, No. 77. Wolffenbuttel. Ceklovský, T., Sabol, M., 2012. Cave bear assemblage from the Sarkanova diera Cave (the Volovské vrchy Mts., Eastern Slovakia). Braunschweiger Naturkundliche Schriften 11, 25e39. Curry, A., 2010. Fate of the cave bear. Smithsonian Magazine on-line version. http:// www.smithsonianmag.com/history-archaeology/Fate-of-the-CaveBear.html? c¼yandpage¼1. Encloe, J.G., David, F., Baryshnikov, G., 2000. Hyenas and hunters: zooarchaeological investigations at Prolom II Cave, Crimea. International Journal of Osteoarchaeology 10, 310e324. Fejfar, O., 1973. Fosilní savci v krasu Ceskoslovenska. Geologický pr uzkum 7, 213e215. Fiebig, M., Pacher, M., 2006. Alpine cave bears and climate in marine isotope stage 3. In: Scientific Annals, vol. 98. School of Geology Aristotle University of Thessaloniki (AUTH), pp. 251e256. a Javorinka vo Vysokých Tatrách. Fudaly, V., Pavlar cík, S., Vadovský, V., 1983. Jaskyn Spravodaj slovenskej speleologickej spolo cnosti 14 (1), 13e20. Grandal d’Anglade, A., 1993. El oso de las cavernas en Galicia: el yacimiento de Cova Eiros. Nova Terra 8, 1e285. Grayson, D.K., Delpech, F., 2003. Ungulates and the Middle-to-Upper Paleolithic transition at Grotte XVI (Dordogne, France). Journal of Archaeological Sciences 30, 1633e1648.
Hain, P.J., 1672. De draconum Carpathicorum cavernis. Miscellanea curiosa medicophisica academiae nature curiosorum. Dec. I. Annus III. Obs. CXCIV. Horacek, M., Frischauf, Ch., Pacher, M., Rabeder, G., 2012. Stable isotopic analyses of cave bear bones from the Conturines Cave (2,800 m, South Tyrol, Italy). Braunschweiger Naturkundliche Schriften 11, 49e54. a v Slovenskej hornatine (Slovenský Janá cik, P., Schmidt, Z., 1965. Medvedia jaskyn raj). Slovenský kras 5, 10e36. Knapp, M., Rohland, N., Weinstock, J., Baryshnikov, G., Sher, A., Nagel, D., Rabeder, G., Pinhasi, R., Schmidt, H.A., Hofreiter, M., 2009. First DNA sequences from Asian cave bear fossils reveal deep divergences and complex phylogeographic patterns. Molecular Ecology 18, 1225e1238. Kosinstev, P., 2007. Late Pleistocene large mammal faunas from the Urals. Quaternary International 160, 112e120. Kostopoulos, D.S., 2006. The Paleoenvironment of the Greek Late NeogeneeQuaternary Ursids. Abstract Book 12th International Cave Bear Symposium, Aridea, Macedonia, Greece, p. 41. Kurtén, B., 1958. Life and death of the Pleistocene cave bear. Acta Zoologica Fennica 95, 1e59. Kurtén, B., 1976. The Cave Bear Story. Life and Death of a Vanished Animal. Columbia University Press, New York, 163 pp. Laughlan, L., Rabeder, G., Sabol, M., 2012. The fossils and taphonomy of the Va zecká Cave e preliminary results. Braunschweiger Naturkundliche Schriften 11, 81e86. Lowe, J.J., Rasmussen, S.O., Björck, S., Hoek, W.Z., Steffensen, J.P., Walker, M.J.C., Yu, Z., the INTIMATE group, 2008. Synchronisation of palaeoenvironmental events in the North Atlantic region during the last termination: a revised protocol recommended by the INTIMATE group. Quaternary Science Review 27 (1e2), 6e17. Marsigli, F., 1726. Danubius Pannonico-Mysicus, observationibus geographicis, astronomicis, hydrographicis, historicis, physicis, perlustratus et in IV tomos digestus, cum tabulis aeri incisis. Hagae, Comitum et Amsterodami. Minieri, M.R., Petronio, C., Sardella, R., Scarano, M., 1995. Le Faune a Mammiferi del Pleistocene superiore dell’Italia peninsulare Padusa. Quaderni 1, 75e87. d z jeskyne Arsamítka. Ceskoslovenský Musil, R., 1953. Jeskynní medve kras 6 (1), 154e156. Axamitky. Anthropozoikum 5, Musil, R., 1956. Nové osteologické nálezy z jeskyne 47e54. Musil, R., . Ursus spelaeus e der Höhlenbär, III. Teil. Weimarer Monographien zur Urund Frühgeschichte 2, 1e112. Musil, R., 1986. Paleobiography of terrestrial communities in Europe during the Last Glacial. Sborník Národního muzea v Praze e Acta Musei Nationalis Pragae, XLI B 1985 (1e2), 1e89. Nordmann, A.V., 1858e1860. Paläontologie Südrusslands I. Ursus spelaeus (odessanus). Helsingfors. Pacher, M., Stuart, A.J., 2009. Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus). Boreas 38 (2), 189e206. a Javorinka vo Vysokých Tatrách (þ271, 49 m). Spravodaj Plu cinský, L., 1996. Jaskyn slovenskej speleologickej spolo cnosti 27 (2), 4e11. 2011. Nové objavy v Javorinke. Spravodaj Slovenskej speleologickej Plu cinský, L., spolo cnosti 42 (2), 1e5. v jaskyni Psie diery. SlovPomorský, F., 1993. Nález zvyskov jaskynného medveda enský kras XXXI, 109e111. Rabeder, G., 1999. Die Evolution des Höhlenbärengebisses. Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, vol. 11, pp. 1e102. Rabeder, G., Nagel, D., Pacher, M., 2000. Der Höhlenbär. Jan Thorbecke Verlag, Stuttgart, 111 pp. Rabeder, G., Frischauf, Ch., Withalm, G., 2006. Conturines Cave and the Ladinic Bear. 32 pp. Sabol, M., 2001. Geographical distribution of Cave Bears (Ursus spelaeus Rosenmüller et Heinroth, 1794) in the territory of Slovakia. Beiträge zur Paläontologie 26, 133e137. Sabol, M., Bendik, A., Stuller, F., Novomeský, F., Necas, L., 2008a. A record of a threelegged cave bear female from the Cave of Izabela Textorisova (the Velka Fatra Mountains, northern Slovakia). Stalactite 58 (2), 31e34. Sabol, M., Döppes, D., Pacher, M., Rabeder, G., Withalm, G., 2008b. Cave bears from the Medvedia jaskyna in the Slovensky raj Mountains (Slovakia): preliminary results. Stalactite 58 (2), 74e77. Schmidt, Z., Chrapan, J., 1970. Datovanie osteologického materiálu fosílnych Ursidae metódou 14C z Medvedej jaskyne v Slovenskom raji. Slovenský kras VIII, 69e82. Stiller, M., Baryshnikov, G., Bocherens, H., Grandal d’Anglade, A., Hilpert, B., Münzel, S.C., Pinhasi, R., Rabeder, G., Rosendahl, W., Trinkaus, E., Hofreiter, M., Knapp, M., 2010. Withering away e 25,000 years of genetic decline preceded cave bear extinction. Molecular Biology and Evolution 27 (5), 975e978. Stiner, M., 1998. Mortality analysis of Pleistocene bears and its paleoanthropological relevance. Journal of Human Evolution 34, 303e326. Bendík, A., Sabol, M., Ne Stuller, F., Novomeský, F., Kraj covi c, J., Straka, L., cas, L., Strecha, J., 2011. The interdisciplinary cooperation of forensic medicine, clinical medicine and palaeozoology: a case of cave bears (Ursus spelaeus) bones. Patologie a soudní lékarství 54 (1), 39e41. Townson, R., 1797. Travels in Hungary with a short account of Vienna in the year 1793 illustrated with a map and sixteen other copperplates. London. Vla ciky, M., 2009. Carnivores from Tren cianske Bohuslavice e Pod Tureckom and Moravany e Lopata II, two Gravettian open-air sites in Slovakia. In: Sabol, M., ovská, Z., Vl Visn cek, L. (Eds.), European Cave Bear Research. Slovenský kras 47 (Suppl. 1), 113e124. Liptovský Mikulás.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
Contents lists available at SciVerse ScienceDirect
Quaternary International journal homepage: www.elsevier.com/locate/quaint
Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians Martin Sabol a, *, Andrej Bendík b, Michal Grivalský c, Martin Grivalský c, Ján Lizák c, Igor Michlík c a b c
Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, SK-84215 Bratislava, Slovakia Slovak National Museum in Martin e Museum of Andrej Kmet, A. Kmet Street 20, SK-03601 Martin, Slovakia Spisská Belá Speleological Group, kpt. Nálepku Street 23, SK-059 01 Spisská Belá, Slovakia
a r t i c l e i n f o
a b s t r a c t New research of the Izabela Textorisová Cave in the Velká Fatra Mts. and the Javorinka Cave in the Vysoké Tatry Mts. yielded important data on Western Carpathians cave bear palaeopopulations. The radiometric dating of cave bear fossils from the Javorinka Cave (c. 51,000 14C BP) represents one of the oldest documented record of cave bears in Slovakia. The site is also the highest documented occurrence of cave bears with an altitude more than 1500 m. The radiocarbon dates from the Izabela Textorisová Cave (30,181 e30,190 cal. BP) represent the last known occurrence of cave bears in the Slovakian Western Carpathians. Ó 2013 Elsevier Ltd and INQUA. All rights reserved.
Article history: Available online xxx
1. Introduction The fossil remains of cave bears (Ursus ex gr. spelaeus) from the Slovak territory are known at least since the Middle Ages. The first mentions of these “dragon bones”, however, occur in historical documents of the seventeenth and eighteenth century (Hain, 1672; Marsigli, 1726; Bell, 1736; Brückmann, 1739). The English traveller Townson (1797) was the first who distinguished the bear fossils among “bones of dragons” from the Demänová Caves. The beginning of more scientific research of cave bear fossil record from Slovak caves can be dated from that time. The first detailed morphological and metrical descriptions of cave bear fossils from the territory under study were published after World War II (Musil, 1953, 1956; Janácik and Schmidt, 1965). To date, fossil remains of cave bears are known from at least 60 Slovak caves (Sabol, 2001), although this number could be higher than 100 sites. With international scientific cooperation with Vienna University and the Austrian Academy of Science, a modern systematic research of some important sites started in 2007 (Sabol et al., 2008b; Laughlan et al., 2012), accompanied also by AMS dating of cave bear fossils (Ábelová and Sabol, 2009). Simultaneously, exploration of other regions expected to have a new cave bear fossil record have been undertaken in highland and alpine
* Corresponding author. E-mail addresses: [email protected] (A. Bendík).
(M.
Sabol),
[email protected]
areas of the Velká Fatra Mts. (Bendík, 2005) and the Vysoké Tatry Mts. (Plu cinský, 2011). Subsequent preliminary study and dating of osteological remains from two sites (Izabela Textorisová Cave in the Velká Fatra Mts. and Javorinka Cave in the Vysoké Tatry Mts.) has produced interesting data on the last occurrence and the geographical distribution of cave bears in the Slovakian Western Carpathians. 2. Localities The Izabela Textorisová Cave is situated in the Konský dol Vale within the Gaderská Valley in the Tlstá National Natural Reservation, Velká Fatra Mts., northern Slovakia (Fig. 1). The site is formed by the Gader limestone (dark-grey compact limestone and lightgrey crinoid limestone, Upper Anisian, Middle Triassic) of the Choc Nappe (Hronikum Unit). The cave consists of a large portal entrance and three halls, more or less connected by horizontal corridors. The bottom of the halls is mostly covered by coarse debris with loam or with layers of guano (thickness to 20 cm). The main location of the discovery of cave bear fossils (named “Hrobka”, Tomb) is situated at the end of one tectonically steep inclined ” (Hall). This corridor is closed corridor in the cave section of “Sien by a cave-in (or collapse), which probably obstructed the original access path for bears. The sedimentary sequence in cave section “Hrobka” consists of an overlying gravel accumulation (covered by 2 cm thick layer of white flowstone), a loamy layer with larger clasts (from 2 cm to 8 cm) and scattered rocks (up to 40 cm in size), and with a thick layer of light-grey clay underlying. The bear fossils have
1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.05.022
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
2
M. Sabol et al. / Quaternary International xxx (2013) 1e8
Fig. 1. Location of the Izabela Textorisová Cave in the Velká Fatra Mts., northern Slovakia.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
M. Sabol et al. / Quaternary International xxx (2013) 1e8
3
at an altitude of 1559 m, with height 7e8 m and width up to 5 m, oriented parallel to the Kolová Valley (Plu cinský, 2011).
been found mainly at depths from 30 to 80 cm above a wet clayey layer. At the same depth, more or less complete skeletons of four to nine cave bear specimens have also been discovered in situ position (Sabol et al., 2008a). ”, a Cave bear remains have also been found in the “Sien 200 300 150 cm section situated at the right side behind the cave entrance. Its base consists of up to 5 cm large clasts and rare large rock blocks mixed with unstratified layers of cave loam, partly with a speleothem. Occasionally, faintly rounded pebbles also occur here, indicating a sporadic underground river. A cave bear record from this cave section (to date, more than 500 fossil remains) consists mainly of teeth, phalanges, and vertebrae. The limb bones are represented in smaller quantities, together with rare skulls and mandible fragments. Some bones show gnawing marks (caused by hyenas?), or fractures (caused by humans?). Javorinka Cave (Fig. 2) is situated in the western part of the Kolový Úplaz Massif (1783 m) on a western slope of the central Javorová Valley on the northern side of the Vysoké Tatry Mts., close to 1254 m elevation (Fudaly et al., 1983; Plu cinský, 1996). The entrance of this corrosive-erosional cave, discovered in 1973 (Fudaly et al., 1983), is located at an altitude of 1218 m (Plu cinský, 1996). The cave is a part of the JavorinkaeNádejná cave system with a current length of 10,501 m (Plucinský, 2011) and consists of carbonate rocks of the Tatrikum Unit (Lower Triassic dark-grey to dark-red sandy laminated clayed shale, Middle Triassic grey dolomite and Guttenstein limestone, light-grey dolomite and rose dolomite with intercalations of hematite, Middle Jurassic red crinoid limestone and grey-rose to grey clayey limestone, and Malm-Neocomian limestone) with tectonic faults oriented NWeSE and NEeSW (Fudaly et al., 1983). Most horizontal and inclined cave
3. Fossil material Fossils of cave bears from the Izabela Textorisová Cave were discovered in 2004 and excavated gradually up to 2007 (Bendík, 2005, 2007; Bendík and Sabol, 2007). More than 1500 bear remains have been found, and are housed in the Slovak National Museum e Museum of Andrej Kmet in Martin town. The fossil record from the site is now under study, and only data on some pathological phenomena (Sabol et al., 2008a; Stuller et al., 2011) with a first attempt of AMS dating (Bendík and Sabol, 2007) are published. To obtain more data on the radiocarbon age of fossils from the cave, some metapodial bones from the “Hrobka” and the ” cave parts were submitted to the VERA Laboratory of Vienna “Sien University. Two of these samples (Mc IV dext. and Mt V sin., described below) yielded numerical dates for the latest occurrence of cave bears in the Slovak territory of the Western Carpathians. Mc IV dext. (HJ/1156) from “Hrobka” cave part is a juvenile or subadult(?) specimen. The grey-white bone without the unjoined distal epiphysis was damaged faintly on its surface and shows no pathological phenomena or marks of post-mortem agents. The ” was adult yellowebrown to brown Mt V sin. (HJ/S-209) from “Sien also damaged on the surface and without pathological phenomena or marks of post-mortem agents (Fig. 3e,f). The measurements of both metapodials are given in Table 1. The fossil record of cave bears from the site is preliminarily assigned to Ursus ex gr. spelaeus. Isolated remains of wolf (Canis lupus) and cervids (cf. Capreolus capreolus, Cervus sp.) have also been found.
Table 1 Measurements (in mm) of cave bear metapodial bones from both studied Slovak caves. Metacarpalia
Maximum length Max. transversal diameter of the proximal epiphysis Max. dorso-plantar diameter of the proximal epiphysis Minimum transversal diameter of the diaphysis Minimum dorso-plantar diameter of the diaphysis Max. transversal diameter of the distal epiphysis Max. dorso-plantar diameter of the distal epiphysis Width of the distal head Metatarsalia
Maximum length Max. transversal diameter of the proximal epiphysis Max. dorso-plantar diameter of the proximal epiphysis Minimum transversal diameter of the diaphysis Minimum dorso-plantar diameter of the diaphysis Max. transversal diameter of the distal epiphysis Max. dorso-plantar diameter of the distal epiphysis Width of the distal head
Izabela Textorisová C. e Hrobka
Javorinka Cave
Mc IV dext. (HJ/1156)
Mc V dext. (HJ/1485)
Mc I dext.
Mc II dext.
Mc II sin.
Mc III sin.
Mc V dext.
e 21.3 31.0 16.4 14.2 e e e
75.5 26.0 28.2 16.2 13.65 23.4 e 15.9
57.5 22.05 16.39 12.14 9.91 19.15 17.0 18.01
70.93 19.09 24.97 17.42 12.41 25.07 18.71 19.55
e 16.52 22.68 14.63 11.1 e e e
80.76 22.54 32.05 16.63 15.21 26.83 22.94 21.91
78.55 28.41 30.99 18.99 16.99 28.19 19.19 25.98
Izabela Textorisová Cave e Hrobka
Izabela Textorisová e Sien
Javorinka Cave
Mt IV sin.
Mt V sin.
Mt III sin.
Mt IV dext.
Mt V sin.
Mt III sin.
(HJ/1345)
(HJ/1343)
(HJ/S-361)
(HJ/S-208)
(HJ/S-209)
79.1 20.2 30.1 17.6 13.2 19.6 e 14.6
92.4 22.8 35.5 18.0 17.5 27.0 e 21.9
83.0 e 31.2 16.5 14.5 24.9 e 19.6
e 19.4 18.4 14.1 12.2 e e e
e 23.4 26.9 11.2 13.0 e e e
corridors formed by river activity, and these are characterized by the abundant occurrence of granitoids, especially in the lower ones. This granitoid material represents alluvial deposits of Kolový Creek, deposited during the Pleistocene gradual incision of the Kolová Valley. In older corridors, the gravel deposits are covered by sandy loam, limestone debris, or thin flowstone. Cave bear fossils have been found in two cave parts: “Medvedí dóm” (Bear Dome) and the “Medvedia chodba” (Bear Corridor). Whereas the “Medvedí dóm” represents a relatively large, 30 m high cave space at an altitude of 1525 m, the “Medvedia chodba” is a 150 m long horizontal corridor
(HJ/1344) 83.2 28.4 29.5 14.2 15.6 25.7 18.2 22.4
84.04 21.79 36.59 16.52 12.65 22.89 e 20.28
Mt IV sin.
Mt V sin.
81.61 21.64 29.35 16.25 13.22 23.19 17.6 20.36
83.66 27.48 30.28 12.74 13.47 24.69 16.86 21.59
Cave bear fossils from the Javorinka Cave were discovered during speleological exploration in 2011 (in the “Medvedí dóm”) and 2012 (in the “Medvedia chodba”) directly on the surface of the cave base (Fig. 4). During sampling of cave bear fossils in the “Medvedí dóm” in summer 2011, a fragment of right maxilla with dentition (C e M2 dext.), I3 dext., p4 dext., m1 dext., and m2 sin. together with metapodials (Mc IeII and Mc V dext., Mc IIeIII sin., Mt IIIeV sin. (of one specimen), and two fragments) and four phalanges were collected. Two groups of bones (metapodials and phalanges) were sent to Vienna VERA-Laboratory for AMS dating.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
4
M. Sabol et al. / Quaternary International xxx (2013) 1e8
The maxilla fragment without P3 (Fig. 3a) is partly covered by a filmy speleothem layer and faintly damaged. The light crowns of preserved cheek teeth are unworn (Stiner’s wear stage III; Stiner, 1998), indicating a young specimen, probably male on the basis of the canine size. The P4 crown morphology resembles Rabeder’s (1999) morphotype F with tiny accessory morphological elements on inner sides of both the metacone and the protocone as well as between the protocone and the paracone. The crown morphology of M1 with the cingulum terminated under the hypocone fully corresponds with Rabeder’s morphotype C3. The M2 crown, with the cusp-shaped cingulum extended as far as the posthypocone, also shows an advanced stage of morphology in the parastyle complex (C3), the mesostyle complex (C), the talon (C/D) as well as in the paracone and protocone inner side morphology (C and 2). Crowns of isolated teeth are also unworn (juvenile specimens) and broken roots of these are partly rounded by water. The morphology of p4 crown (Fig. 3b) indicates Rabeder’s morphotype E with the distinct metalophid, but without conspicuous hypoconid and entoconid. The m1 crown (Fig. 3c) with two mesial metastylids, 4-cusped entoconid complex (B4), 2-cusped enthypoconid, and the distinct hypoconulid separated from the entoconid by one tiny
Brownish metapodial bones of adults (Table 1) are mostly undamaged, and show no pathologies and traces of post-mortem agents (apart of Mc II sin. with probably bitten-off distal epiphysis). Apart from cave bear remains (preliminarily attributed to Ursus ex gr. spelaeus), fossils of Gulo gulo and Martes martes have also been found in the “Medvedí dóm”. 4. Results Uncalibrated radiocarbon dates are given as 14C BP and calendar dates as cal. BP (calculated with OxCal3.10 and the INTCAL09 calibration curve). The first radiocarbon dating of a cave bear left radius from the Izabela Textorisová Cave yielded >45,000 14C BP (Bendík and Sabol, 2007). For that reason, some new samples of cave bear bones were dated by AMS in the VERA Laboratory in Vienna (Table 3). Surprisingly, two new dates demonstrate that the Izabela Textorisová Cave was inhabited by cave bears before c. 24,000 14C BP (c. 30,000 cal. BP). This indicates one of the latest occurrence of cave bears in the Slovakian territory of the Western Carpathians. Thus, the last palaeopopulations became extinct here before the beginning of the Last Glacial Maximum (LGM, c. 30,000 years ago).
Table 3 Radiometric dates of cave bear bone samples from both studied Slovak caves by 14C-AMS in the VERA-Laboratory, Vienna (Austria). Data were calculated with OxCal3.10 and the INTCAL09 calibration curve and correspond to the 2s-confidence level. Probability of the individual time periods is in brackets. For calibration, the pMC (percent Modern Carbon) input option was used (corresponding to a symmetric uncertainty of the uncalibrated 14C age). Site
Lab.-Nr.
Sample name
d13Ca [&]
Uncal. BP age range (2s)
cal. BP age range (2s)
Izabela Textorisová Cave Izabela Textorisová Cave Izabela Textorisová Cave Javorinka Cave
VERA-4041 VERA-5679 VERA-5680 VERA-5682
Radius sin. Mc IV dext. (HJ-1156) Mt V sin. (HJ/S-209) Metapodials
19.5 23.4 23.0 20.7
>45,000 24,640 170 26,460 180 51,000 þ 4500/2900
30,181 cal. BP (95.4%) 29,020 cal. BP 30,190 cal. BP (95.4%) 28,970 cal. BP
a
1.2 0.6 5.5 0.9
1s-error.
accessory cusp represents more advanced morphological stage. Morphology of the m2 crown (Fig. 3d) is relatively advanced, showing a higher level of evolution in the metalophid and metastylid complex, bifurcated mesolophid, 4-cusped entoconid complex, or in the enthypoconid complex (C/D). The measurements of all teeth are given in Table 2.
Table 2 Measurements (in mm) of cave bear teeth from Javorinka Cave in the Vysoké Tatry Mts. L e maximum crown length (antero-posterior dimension), TAW e talonid width, TRW e trigonid width, W e maximum crown width (transverse dimension), WA e anterior crown width, WP e posterior crown width. I3 dext.
L W
15.06 14.87
C1 dext.
L W
25.85 21.49
P4 dext.
L W
21.54 15.28
M1 dext.
L WA WP
29.59 20.14 20.76
M2 dext.
L WA WP
45.90 24.94 23.14
p4 dext.
L W
15.47 10.42
m1 dext.
L TRW TAW
30.15 11.77 14.03
m2 sin.
L TRW TAW
32.91 19.24 19.98
Dating of cave bear samples (metapodials and phalanges) from Javorinka Cave is close to the limit of 14C-dating (Table 3). The date indicates inhabitation of the site by cave bears at least in the Middle Weichselian before 50,000 14C BP. Fossil remains of cave bears found between 1525 and 1559 m in the Javorinka Cave, however, represent the highest occurrence in Slovakia, and can be regarded as a record of a high alpine cave bear palaeopopulation from early MIS 3.
5. Discussion Cave bears from the spelaeus-group (including at least two lineages e Ursus spelaeus sensu scripto and Ursus ingressus) evolved in Europe more than 100,000 years ago (Curry, 2010). The transition between these and their precursors from the deningeri-group is generally considered to have occurred at around the beginning of the Last Interglacial (Rabeder et al., 2000). They were adapted to a wide range of environments, but also occupied different ecological niches (Knapp et al., 2009), in an extreme continental climate. Their established distribution extends eastwards from northwest Spain (Grandal d’Anglade, 1993) across Central Europe and the Urals (Kosinstev, 2007) up to the Altai (Knapp et al., 2009), and from Belgium and the Hartz region of Germany in the north (Rabeder et al., 2000) as far south as Italy (Minieri et al., 1995) and Greece (Kostopoulos, 2006), and southeast to the Crimea (Nordmann, 1858e1860; Encloe et al., 2000). The most northerly record of cave bears is from the northern Urals (62 N; Knapp et al., 2009) and the highest one is from the Italian Conturines Cave, with its entrance at 2800 m above sea level (Rabeder et al., 2006). This high alpine occurrence could be possible only during a warmer Last Glacial period, as the expansion of glaciers during the LGM would have made most mountain areas uninhabitable (Pacher and Stuart, 2009).
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
5
Fig. 2. Javorinka Cave in the Vysoké Tatry Mts., northern Slovakia. Locations where cave bear fossils have been found are marked by ellipses.
M. Sabol et al. / Quaternary International xxx (2013) 1e8
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
6
M. Sabol et al. / Quaternary International xxx (2013) 1e8
Fig. 3. Analysed cave bear fossils from Javorinka Cave (aed) and Izabela Textorisová Cave (eef). a e Right maxilla fragment with C and P4-M2, b e p4 dext., c e m1 dext., d e m2 sin., e fragment of metacarpal bone Mc IV dext. (HJ/1156), f e metatarsal bone Mt V sin. (HJ/S-209); metapodial bones were used for radiocarbon dating.
Cave bears became extinct in northern Eurasia and elsewhere during the last GlacialeInterglacial cycle (Pacher and Stuart, 2009), approximately 25,000 years ago (Curry, 2010). According to former assumptions (Kurtén, 1958, 1976; Musil, 1981), the cave bear could survive in some suitable areas well beyond the LGM, i.e. into the Late Glacial and Holocene, implying their survival until at least c.
12,000e15,000 14C BP. Pacher and Stuart (2009), however, found no evidence that cave bears survived into the Late Glacial in any region. The uncalibrated age data of cave bear remains younger than 30,000 14C BP are known mainly from Balme à Collomb (France), Bärenloch (Switzerland), Lieglloch, Gamssulzen, Ochsenhalt (Austria), and Potocka zijalka (Slovenia). Based on the calibrated data, only the records from Balme à Collomb and Gamssulzen are younger than 30,000 cal. BP. (Fiebig and Pacher, 2006). Cave bears had disappeared in Alps and adjacent areas by about 24,000 14C BP (c. 27,800 cal. BP) (Pacher and Stuart, 2009). Within Central Europe, this datum coincides fairly closely with the cooling at the onset of Greenland Stadial 3 (GS-3) (c. 27,500 cal. BP) (Lowe et al., 2008), well before the LGM, most likely attributable to a marked deterioration in quantity and quality of available plant food (Pacher and Stuart, 2009). Cave bear extinction could thus be plausibly linked to the cooling climate and vegetational changes in GS-3 and GS-2 (broadly “LGM”) (Pacher and Stuart, 2009). On the other hand, climate changes cannot be solely to blame for this event, although the prolonged cold period (LGM) shortened and eliminated growing seasons and changed plant distributions across Europe (Curry, 2010). Though there is evidence of cave bear hunting by ancient hominids (Neanderthals and/or modern humans), it is not likely to have led to the extinction of these animals (Curry, 2010). Rather, different hibernation strategies of cave and brown bears might made a difference (Stiller et al., 2010). Judging from the relative amounts of cave bear and brown bear remains in European caves, cave bears were more dependent on caves for hibernation (Kurtén, 1976; Rabeder et al., 2000). Both modern humans and Neanderthals would have been strong competitors for these caves, and might have forced cave bears into less suitable sites for hibernation (Grayson and Delpech, 2003), where cave bear cubs were more in danger of killing by predators and/or cave bear males. According to the latest aDNA study (Knapp et al., 2009; Stiller et al., 2010), cave bear palaeopopulations began a long, slow decline 50,000 years ago (i.e. approximately 25,000 years before their extinction), around the Middle-to-Upper Palaeolithic transition and well before the LGM began (Curry, 2010). This slow demise
Fig. 4. Fossil accumulations in the Medvedí dóm part of Javorinka Cave, whence some specimens have been collected for analyses.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
M. Sabol et al. / Quaternary International xxx (2013) 1e8
of cave bears suggests that one or more environmental factors must have subtly increased their mortality and/or decreased their reproduction rate (Stiller et al., 2010), although the range of their geographical distribution was not changed (Musil, 1981, 1986). According to Knapp et al. (2009), the cave bear evolutionary history before their extinction involved a rather complex phylogeographic pattern and a more substantial loss of biological diversity at the end of the Pleistocene than has been previously recognized. In any case, their disappearance corresponds broadly with the withdrawal of many extinct and extant mammal species from large area of Europe around the LGM (Pacher and Stuart, 2009). Within the Slovakian territory of the Western Carpathians, the vertical distribution range of cave bears was from 215 m a.s.l. (the Cave) (Sabol, 2001). The Cin cany Cave) to 1559 m a.s.l. (the Murán fossil record from the latter site, however, is documented only by a find of weathered ursid canine (Fejfar, 1973), with attribution to a cave bear highly questioning. Thus, the fossils of cave bears in the “Medvedí dóm” in the Javorinka Cave represent their highest recorded occurrence in the Slovakian Western Carpathians, 1525e 1559 m a.s.l. Although no systematic research of the site has been realized to date and bear remains were preliminary determined only as Ursus ex gr. spelaeus, the advanced morphology of the molars together with a comparison of the M2’s length in relation to the cave part altitude (cf. Horacek et al., 2012: Fig. 2) indicate attribution to the taxon U. ingressus, the presence of which has been validated in Slovakia during a research of Medvedia Cave in the Slovenský raj Mts. (Sabol et al., 2008b). This high alpine occurrence
7
of bears was only enabled by favourable climatic conditions during a relatively warmer Last Glacial period at the beginning of MIS 3 (Ábelová and Sabol, 2009). The date 51,000 þ 4500/2900 14C BP represents one of the oldest documented record of cave bears in the territory under study. The range of the uncalibrated radiocarbon dates from Slovak cave bear fossils extends from 39,100 þ 1100/1000 14C BP (Ceklovský and Sabol, 2012) to 51,100 þ 4700/2900 14C BP (Sabol et al., 2008b), with unaccepted dates of >15,000 14C BP from Medvedia Cave in the Slovenský raj Mts. (Schmidt and Chrapan, 1970) and 15,490 780 to 17,530 900 14C BP from Psie diery Cave (Pomorský, 1993). New radiometric dates from Izabela Textorisová Cave, however, move the upper level of this range to 24,640 170 14C BP or 26,460 180 14C BP (30,181 cal. BP. and 30,190 cal. BP), approximately coincident with a period between Greenland Stadial 4 and 5. These can be assumed as approximate dates of the last occurrence of cave bears in the Slovakian Western Carpathians, with extinction before the LGM. This is in relatively good agreement with results from the Alps and adjacent areas, where cave bears had disappeared by about 27,000e28,000 cal. BP. Fossils of cave bears are entirely absent in Slovakian Upper Palaeolithic sites (e.g. Vla ciky, 2009), whereas sediments of Neanderthal Middle Palaeolithic sites contain these, together with remains of Ursus arctos. Preliminary results from carbon isotopes (d13C food from 28.6& to 22.6&) indicate that the bears fed on plants in a steppe environment, alternating with meadows and light or dense
Fig. 5. Relations among uncalibrated radiocarbon ages of cave bear fossils from Slovak cave sites, altitude of these caves, and isotopic data (d13C). Although there are so far only a few numerical data, a trend of the cave bear palaeopopulations shift to lower (more forested) altitudes with coming LGM can be assumed (data source: Bendík and Sabol, 2007; Sabol et al., 2008b; Ábelová and Sabol, 2009; Ceklovský and Sabol, 2012; Laughlan et al., 2012; and unpublished data).
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022
8
M. Sabol et al. / Quaternary International xxx (2013) 1e8
taiga-like forests (Ábelová and Sabol, 2009). Based on available data, there is also a possibility to suggest movement of cave bear palaeopopulations from high alpine areas to low-altitude (more forested) habitats during the gradually climatic cooling of the coming LGM (Fig. 5). 6. Conclusions The new results of radiometric dating of cave bear fossils from two Slovakian caves (Izabela Textorisová Cave in the Velká Fatra Mts. and Javorinka Cave in the Vysoké Tatry Mts.) yielded important data on Western Carpathians palaeopopulations of this ursid. The date from Javorinka Cave (51,000 þ 4500/2900 14C BP) represents one of the oldest documented record of cave bears in the territory of Slovakia, and can be correlated with the beginning of MIS 3. The site also represents the highest fossil record of cave bears for the area under study with an altitude between 1525 and 1559 m. The new radiocarbon dates from Izabela Textorisová Cave (30,181 cal. BP. and 30,190 cal. BP) represent the last known occurrence of cave bears in the Slovakian Western Carpathians. Palaeopopulations became extinct before the LGM began, what is in good agreement with data from other Central European regions. Acknowledgements The research for this paper was carried out with financial support from the Slovak Research and Development Agency under contact APVV-0280-07 and the Slovak Grant Agency under contract Vega 1/0396/12. The authors also gratefully acknowledge the helpful comments and suggestions of Dr. Evelyne Crégut-Bonnoure and Dr. Silviu Constantin. Their gratitude goes also to Dr. Norm R. Catto for English language review. References Ábelová, M., Sabol, M., 2009. Attempt on the reconstruction of palaeotemperature and palaeoenvironment in the territory of Slovakia during the Last Glacial based on oxygen and carbon isotopes from tooth enamel and bone collagen of cave ovská, Z., Vl bears. In: Sabol, M., Visn cek, L. (Eds.), European Cave Bear Researches. Slovenský kras 47 (Suppl. 1), 5e18. Liptovský Mikulás. Bell, M., 1736. Notitia Hungariae novae historico geographica, divisa in partes qvatvor, quarum prima, Hungariam Cis-Danubiam; altera, Trans-Danubiam; tertia, Cis-Tibiscanam; quarta, Trans-Tibiscanam. Tomus Secundus, Vienna Austriae. p. 579. jaskynného (Ursus spelaeus Bendík, A., 2005. Nové osteologické nálezy medveda Rosenmüller et Heinroth) vo Velkej Fatre. Zborník Kmetianum X, 226e231. Bendík, A., 2007. Zachovanie kostier medveda jaskynneho (Ursus spelaeus Rosenmüller, 1794) v jaskyni Izabely Textorisovej vo Velkej Fatre. Slovensky kras XLV, 219e230. Bendík, A., Sabol, M., 2007. Cave bears from the Cave of Izabela Textorisová (the Velká Fatra Mts., Slovakia) e a state of the art. Scripta Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis 35, 151e156. Geology. Brückmann, F., 1739. Antra draconum Liptovensis. Epist. itineraria, No. 77. Wolffenbuttel. Ceklovský, T., Sabol, M., 2012. Cave bear assemblage from the Sarkanova diera Cave (the Volovské vrchy Mts., Eastern Slovakia). Braunschweiger Naturkundliche Schriften 11, 25e39. Curry, A., 2010. Fate of the cave bear. Smithsonian Magazine on-line version. http:// www.smithsonianmag.com/history-archaeology/Fate-of-the-CaveBear.html? c¼yandpage¼1. Encloe, J.G., David, F., Baryshnikov, G., 2000. Hyenas and hunters: zooarchaeological investigations at Prolom II Cave, Crimea. International Journal of Osteoarchaeology 10, 310e324. Fejfar, O., 1973. Fosilní savci v krasu Ceskoslovenska. Geologický pr uzkum 7, 213e215. Fiebig, M., Pacher, M., 2006. Alpine cave bears and climate in marine isotope stage 3. In: Scientific Annals, vol. 98. School of Geology Aristotle University of Thessaloniki (AUTH), pp. 251e256. a Javorinka vo Vysokých Tatrách. Fudaly, V., Pavlar cík, S., Vadovský, V., 1983. Jaskyn Spravodaj slovenskej speleologickej spolo cnosti 14 (1), 13e20. Grandal d’Anglade, A., 1993. El oso de las cavernas en Galicia: el yacimiento de Cova Eiros. Nova Terra 8, 1e285. Grayson, D.K., Delpech, F., 2003. Ungulates and the Middle-to-Upper Paleolithic transition at Grotte XVI (Dordogne, France). Journal of Archaeological Sciences 30, 1633e1648.
Hain, P.J., 1672. De draconum Carpathicorum cavernis. Miscellanea curiosa medicophisica academiae nature curiosorum. Dec. I. Annus III. Obs. CXCIV. Horacek, M., Frischauf, Ch., Pacher, M., Rabeder, G., 2012. Stable isotopic analyses of cave bear bones from the Conturines Cave (2,800 m, South Tyrol, Italy). Braunschweiger Naturkundliche Schriften 11, 49e54. a v Slovenskej hornatine (Slovenský Janá cik, P., Schmidt, Z., 1965. Medvedia jaskyn raj). Slovenský kras 5, 10e36. Knapp, M., Rohland, N., Weinstock, J., Baryshnikov, G., Sher, A., Nagel, D., Rabeder, G., Pinhasi, R., Schmidt, H.A., Hofreiter, M., 2009. First DNA sequences from Asian cave bear fossils reveal deep divergences and complex phylogeographic patterns. Molecular Ecology 18, 1225e1238. Kosinstev, P., 2007. Late Pleistocene large mammal faunas from the Urals. Quaternary International 160, 112e120. Kostopoulos, D.S., 2006. The Paleoenvironment of the Greek Late NeogeneeQuaternary Ursids. Abstract Book 12th International Cave Bear Symposium, Aridea, Macedonia, Greece, p. 41. Kurtén, B., 1958. Life and death of the Pleistocene cave bear. Acta Zoologica Fennica 95, 1e59. Kurtén, B., 1976. The Cave Bear Story. Life and Death of a Vanished Animal. Columbia University Press, New York, 163 pp. Laughlan, L., Rabeder, G., Sabol, M., 2012. The fossils and taphonomy of the Va zecká Cave e preliminary results. Braunschweiger Naturkundliche Schriften 11, 81e86. Lowe, J.J., Rasmussen, S.O., Björck, S., Hoek, W.Z., Steffensen, J.P., Walker, M.J.C., Yu, Z., the INTIMATE group, 2008. Synchronisation of palaeoenvironmental events in the North Atlantic region during the last termination: a revised protocol recommended by the INTIMATE group. Quaternary Science Review 27 (1e2), 6e17. Marsigli, F., 1726. Danubius Pannonico-Mysicus, observationibus geographicis, astronomicis, hydrographicis, historicis, physicis, perlustratus et in IV tomos digestus, cum tabulis aeri incisis. Hagae, Comitum et Amsterodami. Minieri, M.R., Petronio, C., Sardella, R., Scarano, M., 1995. Le Faune a Mammiferi del Pleistocene superiore dell’Italia peninsulare Padusa. Quaderni 1, 75e87. d z jeskyne Arsamítka. Ceskoslovenský Musil, R., 1953. Jeskynní medve kras 6 (1), 154e156. Axamitky. Anthropozoikum 5, Musil, R., 1956. Nové osteologické nálezy z jeskyne 47e54. Musil, R., . Ursus spelaeus e der Höhlenbär, III. Teil. Weimarer Monographien zur Urund Frühgeschichte 2, 1e112. Musil, R., 1986. Paleobiography of terrestrial communities in Europe during the Last Glacial. Sborník Národního muzea v Praze e Acta Musei Nationalis Pragae, XLI B 1985 (1e2), 1e89. Nordmann, A.V., 1858e1860. Paläontologie Südrusslands I. Ursus spelaeus (odessanus). Helsingfors. Pacher, M., Stuart, A.J., 2009. Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus). Boreas 38 (2), 189e206. a Javorinka vo Vysokých Tatrách (þ271, 49 m). Spravodaj Plu cinský, L., 1996. Jaskyn slovenskej speleologickej spolo cnosti 27 (2), 4e11. 2011. Nové objavy v Javorinke. Spravodaj Slovenskej speleologickej Plu cinský, L., spolo cnosti 42 (2), 1e5. v jaskyni Psie diery. SlovPomorský, F., 1993. Nález zvyskov jaskynného medveda enský kras XXXI, 109e111. Rabeder, G., 1999. Die Evolution des Höhlenbärengebisses. Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, vol. 11, pp. 1e102. Rabeder, G., Nagel, D., Pacher, M., 2000. Der Höhlenbär. Jan Thorbecke Verlag, Stuttgart, 111 pp. Rabeder, G., Frischauf, Ch., Withalm, G., 2006. Conturines Cave and the Ladinic Bear. 32 pp. Sabol, M., 2001. Geographical distribution of Cave Bears (Ursus spelaeus Rosenmüller et Heinroth, 1794) in the territory of Slovakia. Beiträge zur Paläontologie 26, 133e137. Sabol, M., Bendik, A., Stuller, F., Novomeský, F., Necas, L., 2008a. A record of a threelegged cave bear female from the Cave of Izabela Textorisova (the Velka Fatra Mountains, northern Slovakia). Stalactite 58 (2), 31e34. Sabol, M., Döppes, D., Pacher, M., Rabeder, G., Withalm, G., 2008b. Cave bears from the Medvedia jaskyna in the Slovensky raj Mountains (Slovakia): preliminary results. Stalactite 58 (2), 74e77. Schmidt, Z., Chrapan, J., 1970. Datovanie osteologického materiálu fosílnych Ursidae metódou 14C z Medvedej jaskyne v Slovenskom raji. Slovenský kras VIII, 69e82. Stiller, M., Baryshnikov, G., Bocherens, H., Grandal d’Anglade, A., Hilpert, B., Münzel, S.C., Pinhasi, R., Rabeder, G., Rosendahl, W., Trinkaus, E., Hofreiter, M., Knapp, M., 2010. Withering away e 25,000 years of genetic decline preceded cave bear extinction. Molecular Biology and Evolution 27 (5), 975e978. Stiner, M., 1998. Mortality analysis of Pleistocene bears and its paleoanthropological relevance. Journal of Human Evolution 34, 303e326. Bendík, A., Sabol, M., Ne Stuller, F., Novomeský, F., Kraj covi c, J., Straka, L., cas, L., Strecha, J., 2011. The interdisciplinary cooperation of forensic medicine, clinical medicine and palaeozoology: a case of cave bears (Ursus spelaeus) bones. Patologie a soudní lékarství 54 (1), 39e41. Townson, R., 1797. Travels in Hungary with a short account of Vienna in the year 1793 illustrated with a map and sixteen other copperplates. London. Vla ciky, M., 2009. Carnivores from Tren cianske Bohuslavice e Pod Tureckom and Moravany e Lopata II, two Gravettian open-air sites in Slovakia. In: Sabol, M., ovská, Z., Vl Visn cek, L. (Eds.), European Cave Bear Research. Slovenský kras 47 (Suppl. 1), 113e124. Liptovský Mikulás.
Please cite this article in press as: Sabol, M., et al., Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.05.022