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Taphonomische Untersuchungen der Höhlenbärenfundstellen in der Schwabenreith-Höhle bei Lunz am See (Niederösterreich)
... Excavations were conducted at two locations within the cave between 1990 and 1996 (Fladerer, 1992) and in 2015. Of particular interest is excavation site 2, which revealed the following general stratigraphy (from top to bottom; Pacher, 2000;Fig. 2): ...
... Detailed studies of the taphonomy were conducted on 18525 bones and 1796 teeth of cave bear. In total, 16011 of these bones were found at excavation site 2 (Pacher, 2000), where bones of adult bears (65.2%) outnumber those of juvenile (31.4%) and newly born individuals (3.4%). The bear fossils, which belong solely to U. s. eremus (Rabeder et al., 2008), generally lack bite marks. ...
... cave lion) were found in this cave. The skeletons are complete, and there is no evidence of significant post-mortem re-deposition (Pacher, 2000), nor of the presence of humans inside the cave during prehistoric times. Initial attempts to constrain the age of these cave bear sediments included radiocarbon and 230 Th/U dating. ...
The cave bear was a prominent member of the Upper Pleistocene fauna in Eurasia. While breakthroughs were recently achieved with respect to its phylogeny using ancient DNA techniques, it is still challenging to date cave bear fossils beyond the radiocarbon age range. Without an accurate and precise chronological framework, however, key questions regarding the palaeoecology cannot be addressed, such as the extent to which large climate swings during the last glacial affected the habitat and possibly even conditioned the final extinction of this mammal. Key to constraining the age of cave bear fossils older than the lower limit of radiocarbon dating is to date interlayered speleothems using 230Th/U. Here we report new results from one such site in the Eastern European Alps (Schwabenreith Cave), which yielded the highest density of bones of cave bear ( Ursus spelaeus eremus). Although dating of the flowstones overlying this fossiliferous succession was partly compromised by diagenetic alteration, the 230Th/U dates indicate that the bear hibernated in this cave after about 113 ka and before about 109 ka. This time interval coincides with the equivalent of Greenland Stadial 25, suggesting possible climate control on the cave bear's habitat and behaviour.
... Cave bear omnivory was supported by dental mcrowear studies ( Peigné et al. 2009) and the scavenging habit of this species was suggested by tooth marks on bones found at sites where the cave bear was the only carnivoran recorded (e.g. TinTori & ZanalDa 1992, Pacher 2000, PinTo llona & anDrews 2004, PinTo llona et al. 2005, Quiles et al. 2006). However, all these findings can be also interpreted as simply occasional changes in preferential herbivorous habits related to hibernation or seasonal resource availability. ...
... It was suggested based on tooth marks on bones found at sites where the cave bear was the only carnivoran recorded that cave bears showed the scavenging habit (e.g. TinToru & ZanalDa 1992;PaCher 2000;PinTo llona & anDrews 2004;PinTo llona et al. 2005;Quiles et al. 2006). However, no zig-zag HSB so characteristic of typical bone- crushing mammals were observed in U. spelaeus. ...
... Cave bear omnivory was supported by dental mcrowear studies ( Peigné et al. 2009) and the scavenging habit of this species was suggested by tooth marks on bones found at sites where the cave bear was the only carnivoran recorded (e.g. TinTori & ZanalDa 1992, Pacher 2000, PinTo llona & anDrews 2004, PinTo llona et al. 2005, Quiles et al. 2006). However, all these findings can be also interpreted as simply occasional changes in preferential herbivorous habits related to hibernation or seasonal resource availability. ...
... It was suggested based on tooth marks on bones found at sites where the cave bear was the only carnivoran recorded that cave bears showed the scavenging habit (e.g. TinToru & ZanalDa 1992;PaCher 2000;PinTo llona & anDrews 2004;PinTo llona et al. 2005;Quiles et al. 2006). However, no zig-zag HSB so characteristic of typical bone- crushing mammals were observed in U. spelaeus. ...
... The cave has been known locally for a long time but was just investigated only at the end of the 1960s (Hartmann and Hartmann, 1969). Excavations lasted from 1990 to 2000 (Fladerer, 1992;Pacher, 2000). ...
... Fauna: Ursus spelaeus (Pacher, 2000). Site description: The cave is 129 m long. ...
Caves are among the most important sites preserving Quaternary fossils. Owing to the CaCO3-rich environment and the protection against erosion, even remains of early Pleistocene faunas are preserved in caves, while contemporaneous surface deposits have been lost. However, faunal remains cannot be linked to any interglacial or glacial period since no species exists which is characteristic of any specific period. Reliable dating of such remains is therefore required. This is now possible applying 230Th/U dating of speleothems. ESR dating of speleothems or 230Th/U dating of bones is, however, of disputable value. Dating of the base and top speleothem accretional layers permit assigning Pleistocene faunal remains to the MIS chronology. In this paper, we present for the first time an overview of all numerically dated palaeontological cave sites in Central Europe between MIS 5 and MIS 8. From twelve sites, a total of 31 strata were dated, most of them deposited during MIS 5; the rest belongs to MIS 6 and MIS 7; and only one sample representing MIS 8 provided reliable numerical dates. Numerically dated palaeontological cave sites older than MIS 8 are not known.
... Age-scoring and survival analyses for Niedźwiedzia cave bears show a peak mortality rate (27%) between 1.5 and 3 years (stage II), and 17% for newborns (< 1.5 years) and 11% for individuals aged 12-16 years. This aligns with Schwabenreith-Höhle's findings, emphasising stage II prevalence 52 . Unlike Slovenian (DivJe babe I, Mokrica, and Potocka zijalka), Belgian (Goyet), Romanian (Peștera cu Oase), and German (Bärenhöhle) sites with abundant stage I remains, Niedźwiedzia reveals different mortality patterns [53][54][55][56][57][58] . ...
This comprehensive study examines fossil remains from Niedźwiedzia Cave in the Eastern Sudetes, offering detailed insights into the palaeobiology and adversities encountered by the Pleistocene cave bear Ursus spelaeus ingressus. Emphasising habitual cave use for hibernation and a primarily herbivorous diet, the findings attribute mortality to resource scarcity during hibernation and habitat fragmentation amid climate shifts. Taphonomic analysis indicates that the cave was extensively used by successive generations of bears, virtually unexposed to the impact of predators. The study also reveals that alkaline conditions developed in the cave during the post-depositional taphonomic processes. Mortality patterns, notably among juveniles, imply dwindling resources, indicative of environmental instability. Skeletal examination reveals a high incidence of forelimb fractures, indicating risks during activities like digging or confrontations. Palaeopathological evidence unveils vulnerabilities to tuberculosis, abscesses, rickets, and injuries, elucidating mobility challenges. The cave’s silts exhibit a high zinc concentration, potentially derived from successive bear generations consuming zinc-rich plants. This study illuminates the lives of late cave bears, elucidating unique environmental hurdles faced near their species’ end.
... This is also indicated by the activities aimed at the extraction of meat and marrow, the consumption of which is not strictly fixed in a timeframe, as it depends on taste preferences and food tolerances unknown today (Stiner et al., 1996). Given our certitude that the period of procurement has been established to be early spring at Fumane and early spring and/or preceding hibernation at Rio Secco, we suggest, following Stiner's ethnographic observations of Native American Indians (1994) and Pacher (2000) and Münzel and Conard's (2004a, b) analyses of ethnographic data on bear hunting since the 17th century, that the Neanderthal predation model was based on the opportunistic predation of bears during the first stages of hibernation and during winter, the favored season for hunting this plantigrade. It remains, however, problematic to imagine the possibilities and existing hunting techniques on the part of Neanderthal hunters in northern Italy, even knowing that the lithic assemblages are extremely lacking in stone points and that the prey species are among the largest represented of the carnivore order. ...
Cave bear (Ursus spelaeus), brown bear (Ursus arctos), and Neanderthals were potential competitors for environmental resources (shelters and food) in Europe. In order to reinforce this view and contribute to the ongoing debate on late Neanderthal behavior, we present evidence from zooarchaeological and taphonomic analyses of bear bone remains discovered at Rio Secco Cave and Fumane Cave in northeast Italy, an extended geographic area north of the Adriatic Sea. The remains from both caves come from layers dated to 49-42 ky cal. BP, and suggest close interactions between humans and bears, with data not only limited to the association of Mousterian lithic artifacts with numerous bear remains, but also the detection of clearly preserved traces of human modification such as cut and percussion marks, which enable a reconstruction of the main steps of fur recovery and the butchering process. Examples of Neanderthal bear exploitation are extremely sporadic in Europe, and Grotta Rio Secco and Grotta Fumane can be considered rare cases of remain accumulations generated by the human predation of bears of varied age classes during or near the end of hibernation. All of this evidence suggests that bears had a strategic role in the nomadic economy of Neanderthal hunting groups.
... The study of carnivore impact on bones from a cave bear thanatocoenosis allows researchers to: (i) detect a taxon/ species presence in the absence of osteological remains of that species, (ii) determine carnivore patterns of carcass consumption, and (iii) collect valuable data for palaeofaunistic reconstruction. Using a taphonomic approach many European cave bear (Ursus spelaeus) sites have been intensively investigated and the carnivore impact on U. spelaeus remains has been assessed over the past few decades (e.g., Tintori & Zanalda 1992; Stiner et al. 1996; Pacher , 2004 Bona 2003; quilès 2003; DomínguezRodrigo & Piqueras 2003; Döppes 2004; Fosse et al. 2004; Pinto-Llona & Andrews 2004; Martínez-Sánchez et al. 2012; Arilla et al. 2014; Fourvel et al. 2014). Although the Romanian Carpathians have great potential for addressing these issues, relevant publications are lacking, except for Jurcsák et al. (1981), quilès et al. (2006) and Pacher & quilès (2013. ...
In taphonomy, the study of carnivore modification of fossil bones and the analysis of their dispersion represent the best approach to assessing the extent of bone modification and displacement for a given bone assemblage. Here we analyze the excavated bone deposit from Urşilor Cave, a well-documented and fossil-rich Upper Pleistocene cave bear site from the Romanian Carpathian Mountains. More than 1400 limb bones or bone remains were analyzed (NISPleft and right = 1424) and 69 measurable puncture marks were identified, measured and morphologically analyzed. Moreover, for assessing the degree of bone scattering, almost 540 cave bear limb bones and mandibles were refitted and the Index of Skeletal Disjunction (ISD) was calculated for the entire bone assemblage. More than 30 % of the analyzed cave bear limb bones were affected by carnivores: the ulnae were the most affected (39.3 %) while the humeri and femora were less modified (24.7 % and 25.5 %, respectively). The range of variation in size of the puncture marks, the morphological features of various tooth marks and the faunal composition of the studied bone assemblage indicate that at least two carnivore taxa are responsible for the bone modifications. The results obtained for the ISD index indicate higher displacement for femora when compared to other bones (e.g. tibiae, mandibles, humeri). Our analyses of bone modifications caused by carnivores indicate a low level of the scattering of intensely modified (by in situ consumption) bones, and notable carnivore impact on the configuration of the bone assemblage.
Key words: Ursus spelaeus, taphonomy, ISD, tooth marks, Urşilor Cave, Romanian Carpathians.
Vpliv zveri na kosti jamskih medvedov iz jame UrŞilor (SZ Romunija) in analiza njihove razkropljenosti
Proučevanje sledov zveri na fosilnih kosteh in razkropljenosti kosti v prostoru je najboljše tafonomsko orodje za pridobivanje podatkov o obsegu takšnih pojavov znotraj posameznega paleontološkega zbira. V raziskavi obravnavamo kosti jamskega medveda iz jame Urşilor, dobro raziskanega mlajšepleistocenskega najdišča v Romunskih Karpatih. Analizirali smo več kot 1424 dolgih cevastih kosti okončin ali njihovih odlomkov in pri tem prepoznali, izmerili in morfološko analizirali 69 odtiskov zob. Poleg tega smo izračunali indeks razkropljenosti okostja (Index of Skeletal Disjunction, ISD) za celotni paleontološki zbir, zaradi česar smo sestavili skoraj 540 dolgih cevastih kosti okončin in spodnjih čeljustnic jamskega medveda. Sledove zveri smo prepoznali na več kot 30 % kosti okončin, najpogosteje na komolčnicah (39.3 %), nekoliko redkeje pa na nadlahtnicah in stegnenicah (24.7 % oz. 25.5 %). Variabilnost v velikosti odtiskov zob, njihovih morfoloških značilnostih in favnistični sestavi proučevanega paleontološkega zbira pričajo o tem, da sta ugotovljene spremembe medvedjih kosti povzročili vsaj dve različni zveri. Vrednosti indeksa razkropljenosti okostja kažejo na izrazitejšo razkropljenost stegnenic v primerjavi z drugimi kostmi (npr. golenicami, nadlahtnicami, spodnjimi čeljustnicami). Naša analiza kaže na skromno razkropljenost intenzivno obgrizenih kosti (zaradi hranjenja v sami jami) in znaten vpliv zveri na konfiguracijo paleontološkega zbira.
Ključne besede: Ursus spelaeus, tafonomija, ISD, ugrizi, jama Urşilor, Romunski Karpati.
... To cope with this, a calibration curve was established using independent dating methods such as dendrochronology (until ~10,000 BC), Uranium-Thorium dating of speleothems and corals, and varve counting of terrestrial and marine sediments. The limit of the method is due to the fact that after approximately 10 half-lives (half-live of 5,730±40 years) only less than 1 permille of the original 14C remains, hence no material older than 50,000 years can be dated reliably with this method (Reimer et al., 2013;Olsson, 2009). ...
New AMS dating for three Austrian sites were conducted on cave bear bones at the Klaus-Tschira-Archaeometry Center in Mannheim, Germany. In total 14 new dates will be presented. The oldest date is 48 ka BP. The faunal remains from the Schwabenreith Cave, located near Lunz (Lower Austria), only consist of cave bears from the taxa Ursus spelaeus eremus. The basal and top flowstone layers of excavation area 2 yielded U-Th ages of 116±5 ka and 78+30/-23 ka BP, respectively. In the Herdengel Cave, located in the same region, the remains of U. sp. eremus and U. ingressus were found. A basal flowstone layer yielded a U-Th age of 112+12/-11 ka BP. The Brettstein Cave system in the Totes Gebirge (Styria) represents the two cave bear taxa U. sp. eremus and U. ladinicus. Dated cave bear bones were only known to be older than 49 ka BP.
The new AMS dates include six bone remains from Schwabenreith Cave dated in the period from 34 ka to 48 ka BP. New dating results from the Herdengel Cave show a very close timespan from 44 ka to 48 ka BP. And finally the bears of the Brettstein Cave represent one of the youngest dated remains (22.5 ka to 35 ka BP) in the Alps.
... Using this approach, over the last few decades, many European cave bear sites were intensively investigated (e.g. Tintori and Zanalda, 1992;Stiner, 1996;Pacher, 2000Pacher, , 2004cBona, 2003;Quilès, 2003;Döppes, 2004;Pinto-Llona and Andrews, 2004;Fourvel, 2013) and the carnivore impact on U. spelaeus remains has been assessed. For the Romanian Carpathians, the literature concerning this issue is rather scarce (Jurcsák et al., 1981;Quilès et al., 2006;Pacher and Quilès, 2013) although the potential for such studies is tremendous. ...
Cave bears are among the best studied Upper Pleistocene extinct
species due to the high abundance of their fossils in caves across Europe. Ursus minimus (Pliocene) was the common ancestor of the the cave, brown and Asiatic black bears. The speleoid and the arctoid lineages split 1.6 million years ago (Bon et al., 2008) and cave bears became extinct around 27,500 years ago (Pacher and Stuart, 2009). Until recently, the cave bear’s distribution was thought to be restricted to the European continent, but the
study of Knapp et al. (2009) confirmed the existence of cave bears on the Asian continent. Morphological studies based on dental features (Rabeder, 1995) that later were confirmed by molecular results (Hofreiter et al. 2004), proposed two distinct species within the European cave bear group: U. spelaeus and U. ingressus (Rabeder et al., 2004). The first species corresponds to the western clade of the group, while the second species has
an Eastern European distribution. For reasons of consistency and lack of molecular data for the Romanian cave bears, the term U. spelaeus (sensu Rosenmüler and Heinroth, 1794) will be used throughout this study.
... Fauna: U. spelaeus (Pacher, 2000). ...
Caves are terrestrial depositories that preserve a large variety of organic and inorganic remains. These may contain important Quaternary climatic and ecological information. Most of the faunal remains, however, cannot be linked to any Interglacial or Glacial period exclusively. Reliable dating of such remains is therefore required. Experience has, however, shown that ESR dating of speleothems or 230Th/U dating of bones are of disputable value. Only TIMS-230Th/U dating of speleothems appears to yield reliable ages. Dating the bottom and top of speleothem layers permit assigning Pleistocene faunal remains to the OIS chronology if the deposition of the speleothems and the faunal remains are clearly correlated. Care must be taken to consider the depositional situation of each site before interpreting any age dates. In this paper we present an overview of all numerically dated paleontological cave sites in Central Europe between OIS 5 and OIS 8. A total of 25 strata were dated from 13 sites, most of them deposited during OIS 5; the rest belonging to OIS 6 and 7. Numerically dated paleontological sites older than OIS 8 are not known.
... At sites where cave bear is the only member of the Carnivora recorded, a low incidence of tooth marks on the bones suggests occasional scavenging of cave bear remains by other cave bears (Tintori & Zanalda 1992;Pacher 2000a). These activities might reflect the need for food or minerals on emerging from hibernation. ...
The cave bear (Ursus spelaeus) was one of several spectacular megafaunal species that became extinct in northern Eurasia during the late Quaternary. Vast numbers of their remains have been recovered from many cave sites, almost certainly representing animals that died during winter hibernation. On the evidence of skull anatomy and low δ15N values of bone collagen, cave bears appear to have been predominantly vegetarian. The diet probably included substantial high quality herbaceous vegetation. In order to address the reasons for the extinction of the cave bear, we have constructed a chronology using only radiocarbon dates produced directly on cave bear material. The date list is largely drawn from the literature, and as far as possible the dates have been audited (screened) for reliability. We also present new dates from our own research, including results from the Urals. U. spelaeus probably disappeared from the Alps and adjacent areas – currently the only region for which there is fairly good evidence –c. 24 000 radiocarbon years BP (c. 27 800 cal. yr BP), approximately coincident with the start of Greenland Stadial 3 (c. 27 500 cal. yr BP). Climatic cooling and inferred decreased vegetational productivity were probably responsible for its disappearance from this region. We are investigating the possibility that cave bear survived significantly later elsewhere, for example in southern or eastern Europe.
Sex-ratio determination is essential for palaeontological and palaeoethological researches on fossil taxa, especially for cave bear which shows an important sexual dimorphism. “Classical” methods use uni- and bivariate plots to fix an arbitrary limit between sexes. Therefore, a more robust statistical method, termed “mixture analysis”, has been applied in this study to determine the sex-ratio of the cave bear from Schwabenreith, Gamssulzen (Austrian Alps), Fate, Basura and Badalucco (northwest of Italy). This method has been previously calibrated on the post-cranial material of cave bear from Fate (QUILES & MONCHOT 2004). In order to define sexual dimorphism for the whole skeleton, seven elements (lower canine, lower and upper second molars, scapholunar, navicular, metacarpal 3 and metatarsal 3) have been selected for their reliability and their abundance in studied assemblages. A quantified comparison between results of bivariate plots and mixture analysis highlights the better reliability and rigor of the new method. The latter, however, also offers a standardized and replicable procedure which provides a calculated cut-off point and a misclassification error for each dimension, permitting the analyst to determine precise sexual groups as well as identifying those specimens that cannot be sexed. Cheek teeth can be sexed as reliably as lower canine and post-cranial elements, providing statistically equivalent results. Lower canine, second upper molar and scapholunar are the most effective bones for determining sex-ratio in cave bear.
The fossil population structure of the cave bear from the Mokrica cave was evaluated to provide new data concerning the behaviour and mortality of this extinct species. Age at death was estimated for 128 different individuals by analysing cementum increments, root formation and crown wear of left M, teeth. After the frequency distribution of specimens through one-year intervals, the mortality trends can be estimated for various lifetime periods, and interpreted in accordance with data for present-day bears. The original death assemblage was presumably juvenile-dominated. Extremely fragile molars of less than 6 month old cubs did not get preserved. Yearlings are the most numerous age class in the fossil population from the Mokrica cave. Mortality drastically dropped after cave bears survived their first hibernation in the second winter. The lowest mortality rate was observed in the 9-15 years age group, when cave bears would be expected to be in their prime. The oldest age recorded by cementum analysis is approximately 30 years, which indicates that the maximum life span was similar to present-day bears. Study of dental tissues shows that the mortality in the cave was seasonally restricted - the majority of deaths in the cave occured during winter and in early spring. Sex structure of the fossil population has been studied on the sample of 750 canines. The significantly higher proportion of mates in the group of older juveniles and subadults could be explained by the fact that the weaning period is more critical for mates also in present-day bears. In young adults and prime adults the mortality was presumably higher in females. The sex structure of adult bears, especially in the sample of older individuals, indicates that the Mokrica cave was used as winter den mostly by solitary males.
Potočka zijalka is a rich archaeological and palaeontological site. The following paper gives a first overview concerning the taphonomic analyses of the Upper Middle Wuermian cave bear assemblage from this cave. Special emphasis is given to the spatial distribution analyses and the method of re-articulation of cave bear bones. First results indicat e a scattered and sorted distribution of remains and revealed, that articulated finds alone are no evidence for cave bear remains on primary deposition. The taphonomic agents responsible for this preservation pattern at Potočka zijalka are probably cave bears themselves, wolves and non-biological site formation processes. No evidence of human influence on the assemblage was detected so far on the material available for this study.
Potocka zijalka is a unique site with rich archaeological and palaeontological remains. From 1997 onward new excavation campaigns were carried out in co-operation with the Institute of Geology in Ljubljana and the Institute of Palaeontology in Vienna. This paper comprises the state of investigation and gives a short overview of major discussion points concerning the role of Palaeolithic man and cave bears at this site.
Based on tomographic (CT) images the description of some internal details of the mummy of young woman aged 21-23 years was given. The mummy was earlier confirmed to be of Egyptian origin and dated to the Ptolemaic Period. Tomographic images enabled to make an exact copy of the skull and allowed for the reconstruction of facial appearance by Gierasimov’s method. The skull was partially filled with resin, traces of
which were also visible in the sphenoid sinus and maxillary sinuses. Additionally the entire spinal canal (including the lumbar region) was filled with resin, what was especially interesting since such a practice has not been described so far. Lower limbs were reinforced by the placement of the two 20 cm long pins in the knee area. Upper limbs were crossed on the mummy’s chest. A spindle-shaped object about 4.5 cm long was found under the left hand. In CT scans the object appeared to possess a radiodense area in the centre surrounded by delicate substance of lower radiodensity. The object proved to be a bulb of Narcissus
(cf. tazetta L. fam: Amaryllidaceae) that was the only species of Narcissus growing wild in northern Egypt. Small fragments of subfossil Narcissus bulbs served as talismans and were found inside mummies at several localities in Egypt. However, the case of a mummy equipped with a complete bulb has not been described, yet.
For this investigation an amount of 4459 metapodial bones of Ursus deningeri and Ursus spelaeus was treated. Up to eight measurements were taken and four indices were calculated to follow the evolution of metacarpus and metatarsus and the development of the metapodials in particular.
The material was taken from eight different sites in Austria and Southern Tyrol (Italy), differing in age and altitude. These sites are: Hundsheimer Spalte (Lower M–Pleistocene, n < 30), Repolust cave (Upper M–Pleistocene), Schwabenreith cave (Early Wurmian), Herdengel cave (Early – Middle Wurmian), Ramesch–Knochenhöhle (Middle Wurmian, two separated stratigraphic levels, high alpine), Conturines cave (Early – Middle Wurmian, Southern Tyrol, high alpine), Windener Bärenhöhle (Middle Wurmian) and Gamssulzen cave (Late Wurmian, the standard for cave bear faunas, all other faunas will be related to the population of this cave). The following points were considered:
1. Is there an evolution of metapodial bones leading from Ursus deningeri to Ursus spelaeus and if so, is this evolution related to the evolution of the P4/4–Index ?
2. Are there any peculiarities in the material from the high alpine cave bear faunas ? Which position do these bears obtain within the cave bear group ?
3. Is it possible to determine the sex–ratio by means of metapodial bones ?
4. As it became obvious that there is one more species of bear in the Windener Bärenhöhle these findings are described as well.
The Mladeč Caves lie west of the village Mladeč near Olomouc in Moravia. The cave system developed in the Devonian limestone of the Třesín Hill, and was discovered in the course of limestone quarrying. Following Svoboda (this volume), four sites are distinguished inside and above the hill. Site I is situated in a large cave consisting of several halls, collapsed chimneys, and corridors. Older names of this site include "Fürst-Johanns-Höhle" and "Bočkova díra". After its discovery in 1826 or 1829, finds of a "giant" and of animal bones were reported from Hall A behind the entrance (Szombathy, 1925, 4; Maška, 1886). Josef Szombathy, from the Naturhistorisches Museum in Vienna, carried out the first scientific excavations in 1881 and 1882 in Hall D (Szombathy, 1882; Hochstetter, 1883). In 1902, additional parts of the cave were opened and finds of animal bones were determined by Knies (Szombathy, 1904). The exact locality of this survey is not known. At about the same time, a quarrel over the claim of ownership occurred. In April 1902, the owner of the "Plavatisko", the area of the Třesín Hill directly above the cave, closed the entrance to the Mladeč Cave and opened up a new access directly from his ground (after Smyčka in Szombathy, 1904). The year 1902 is inscribed on the rock wall in the middle of the chimney leading into Hall A (Svoboda, 2000, 527). Thus, it seems likely that the chimney into A was re-opened during the dispute. From 1903 onwards, Knies examined different localities in Halls D and E, probably around the large debris cone that he first observed (Svoboda, 2000, 530; Szombathy, 1925, 9). At that time, the first arrangements to accommodate public visits inside the cave had already begun (Svoboda, 2000, 530). In 1911, the Krajínski musejní spolecnost v Litovli (Museum Society in Litovel) became the owner of the total area of the cave (Oliva, 1989, 53). In order to better adapt the cave for public visits, large-scale earth removals took place. Some parts of the cave were levelled of up to 3 or 4 meters and as a result, deeper parts of the cave were discovered (Szombathy, 1925, 1, 9). A few juvenile bear remains were mentioned only from the Netopiře jeskyně (bat cave) southeast of Hall E. The corridor "s" served as a second entrance after removal of the filled in sediment. Szombathy (1925, 9) assumed here an original horizontal entrance to Mladeč Cave I, but during the Late Pleistocene this corridor was already filled by earlier, perhaps Middle Pleistocene sediments (Svoboda, 2000, 534). Fürst and Smyčka carried out additional important excavations in 1922. The exact position of their survey is not known but it is assumed to be near findspot "e" (Oliva, 1989, 53). Szombathy (1925, 10) speaks of two separate "fireplaces" with animal bones, human remains, and bone artifacts in the surrounding area. Spot "II" on the ground map of Jelínek (1983) might indicate one of these localities (see Fig. 3b). Northwest of the "fireplaces", mainly bovid remains are mentioned. This locality is the continuation of Szombathy's findspot "d". More recent investigations in Hall D were organised by Jelínek from 1958 to 1963, and concerned mainly Middle Pleistocene layers (Svoboda, 2000, 532; Jelínek, 1983, 1987). Mladeč Cave II is located about 50 steps west of the present entrance to site I. It was discovered and subsequently destroyed in 1904 during quarrying operations. A week later, Knies examined the site and collected various finds (Szombathy, 1904; Svoboda, 2000, 531). A third site, "Podkova" cave, lies on the northern slope of the hill, and is separated from the main cave system. Directly above the karstic system lies site IV, called "Plavatisko". Here a Gravettian open-air site was located, but the Třesín Hill was settled throughout prehistory (Jelínek, 1983). For a more detailed description of the sites and the history of investigation at the Mladeč Caves, see Svoboda (this volume) and Svoboda (2000). Sites I and II are famous mainly for their rich anthropological material. More than 100 specimens of modern humans are reported (see Svoboda et al., 2002, 957). Based on skulls and maxillae, the preserved material represents seven or eight individuals, including one child (see list in Jelínek, 1983). Immediately after the discovery of the human bones, their Pleistocene age was questioned (Maška, 1886). The alleged contemporaneousness of reindeer and human remains as emphasized by Hochstetter (1883), probably favoured a Magdalenian age of the assemblage but this was already doubted by Hoernes (1903). Finally, Bayer (1922) placed the remains in the Aurignacian horizon. He summarized various arguments to prove his assumption. Among them was evidence of cave bears, one perforated bear tooth, and above all, bone points with a massive base. Bayer (1922) regarded these artifacts as diagnostic of the Early Upper Paleolithic, and introduced the Mladeč Caves as an eponymous site for bone points with a massive base. The various findspots in the Mladeč Caves have produced about 40 bone points but only a few stone artifacts. According to Svoboda (2000, 531), none of the lithic artifacts are diagnostic. Following Oliva (1989, 54), only one of the preserved specimens found at today's entrance area can be ascribed to an Aurignacian tradition. In addition, Valoch (1995, 73) and Jelínek (1983) report an artifact found in corridor "c", which could be of Middle Pleistocene age (Svoboda, 2000, 531). The rich faunal material consists of large mammals and small vertebrate remains. Various authors have published different faunal lists (see summary in Musil, 2002), but the material has not been examined in detail until today.
The cave bear spread from Western Europe to the Near East during the Riss glaciation (250 KYA) before becoming extinct approximately 12 KYA. During that period, the climatic conditions were highly dynamic, oscillating between glacial and temperate episodes. Such events have constrained the geographic repartition of species, the movements of populations and shaped their genetic diversity. We retrieved and analyzed ancient DNA from 21 samples from five European caves ranging from 40 to 130 KYA. Combined with available data, our data set accounts for a total of 41 sequences of cave bear, coming from 18 European caves. We distinguish four haplogroups at the level of the mitochondrial DNA control region. The large population size of cave bear could account for the maintenance of such polymorphism. Extensive gene flow seems to have connected European populations because two haplogroups cover wide geographic areas. Furthermore, the extensive sampling of the deposits of the Scladina cave located in Belgium allowed us to correlate changes in climatic conditions with the intrapopulational genetic diversity over 90 KY.
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