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Extinction pattern of Alpine cave bears - new data and climatological interpretation
Historical Biology
Abstract
Cave bears have disappeared from the Alps from different altitudes at different times. The temporal progression of the HDEL (Height Dependent Extinction Line) – a compilation of the geologically most recent radiocarbon dates per altitude level – is not consistent with the general cooling of the temperatures from about 45 ka BP. The cave bear sites of the Northern Alps with the most recent radiocarbon ages are not situated in the lowlands but in caves in altitudes of 1,500 m to 1,700 m above sea level (a.s.l.).
Cave bears fed almost exclusively on herbs and leaves. It was assumed that with the general cooling in the OIS 3 since about 45 ka BP also the migration of the alpine elements into the lowlands took place. It could be recognized that the populations in the lower situated cave bear site became earlier extinct than the cave bear population in the higher altitudes.
With new radiocarbon dates, done at the Curt-Engelhorn-Center Archaeometry at the Reiss-Engelhorn-Museen in Mannheim (Germany), the HDEL can be determined much more precisely and the causes of gradual extinction are also better understood.
... However, cave bear bones and teeth are not only found in caves, as the name suggests, but also at open air sites in the lowlands, such as Bobenheim-Roxheim in the Upper Rhine Graben. After determining the distinct species of the bear remains, usually radiocarbon dating is applied (Döppes et al., 2016;Rossi et al., 2018;Döppes et al., 2019). When dealing with radiocarbon ages, a closer look at the radiocarbon section of this paper will be helpful as this provides hints on how radiocarbon data should be presented or interpreted. ...
... In the following paragraphs, we discuss the outcomes of different analytical methods, ways of data presentation and combining them. For comparison, we include stable isotope data from the literature that are uncorrected regarding altitude or bone/ tooth type (Kneussl, 1972;Kneussl, 1973;Blant et al., 2010;Bocherens et al., 2011;Münzel et al., 2011;Spötl et al., 2014;Döppes et al., 2016, Krajcarz et al., 2016Döppes et al., 2019). ...
... To monitor a possible dependency of the δ 15 N data on altitude, Fig. 6: Stable isotope data of cave bear bones from this study (larger symbols) in comparison to data from other publications on European cave bears (small symbols) (Blant et al., 2010;Döppes et al., 2016;Döppes et al., 2019;Kneussl, 1972Kneussl, , 1973Krajcarz et al., 2016;Münzel et al., 2011;Spötl et al., 2014 the data of this study were plotted with those from other studies ( Fig. 7). Values of less than 0 ‰ are regarded as unusually low. ...
Radiocarbon dating is generally the first choice to date younger Upper Pleistocene organic material. In caves, skeletal remains of animals or humans are often the only preserved datable finds. Routinely, collagen is extracted from bones for radiocarbon dating as it does not readily exchange carbon with its surrounding. However, collagen can also deteriorate and is sometimes not at all or only insufficiently preserved. Apart from the age determination, dietary habits are of special interest and are reconstructed using the stable isotopes of carbon (δ13C) and nitrogen (δ15N). We describe the major challenges of radiocarbon dating and stable isotope analysis of cave bear remains from the Austrian Alps, Franconia (Germany) and fossil finds at a site near Bobenheim-Roxheim (Germany), but also the informative potential of successful application of the methods. We intent to sharpen the knowledge what to look out for to be able to obtain reliable ages of cave bear and other bone material. Moreover, we present two additional scientific methods, luminescence dating and dendroecology as further possibilities to disclose information about sites, their chronology and climatic conditions.
... ed these regions far above 2,000 meter altitude. Not only from the negative correlation of the 13 C levels but also from the extinction pattern of the cave bears in the Northern Alps (DÖPPES et al., 2018), one can conclude that dryness increased over time during the cave bear era. pretože vysokohorské medvede jaskynné obývali tieto oblasti vysoko nad 2 000 metrov nad morom. ...
... An age of around 44,000 years is combined with an "older than" date, as in numerous alpine bear caves (DÖPPES et al., 2018). The cave bear era began in the Carpathians already 50,000 years ago calBP and lasted until at least 45,000 years ago. ...
... Vek približne 44 000 rokov je kombinovaný s údajom "starším ako", ako je to aj pri mnohých vzorkách medveďov jaskynných z alpského prostredia (DÖPPES et al., 2018). Éra medveďov jaskynných sa začala v Karpatoch už pred 50 000 rokmi calBP a trvala do obdobia najmenej pred 45 000 rokmi. ...
... 5) and thus lie within the time range of the other »plateau bear« ages determined so far (Fig. 6). The period during which the »plateau bears« inhabited the karst plateaus of the Northern Calcareous Alps corresponds to almost the entire Middle Wurmian, also called the »Alpine Cave Bear Era« (Döppes et al., 2018). ...
... Symbole: Rote Punkte und Linien bezeichnen die Mittelwerte und Fehlergrenzen der molekularen Datierung, schwarze Punkte und Linien bezeichnen die Mittelwerte und Fehlergrenzen (2σ _calBP) der Radiokohlenstoffdatierung nach der AMS-Methode. Daten nachDöppes et al., 2018;2023;Frischauf et al., 2022;Kavcik-Graumann et al., 2022, Rabeder et al., 2019 Withalm et al., 2022 und dieser Artikel (Tab. 5). ...
The fossil fauna of the Schottloch was excavated and described 140 years ago. A modern analysis of the extensive material has now been carried out for the first time. The taxonomic position of the cave bear remains, the degree of mountain adaptation compared to other high alpine cave faunas and the geological period during which the cave bears used the Schottloch as wintering quarters is the main research objective of this study.
... 5) and thus lie within the time range of the other »plateau bear« ages determined so far (Fig. 6). The period during which the »plateau bears« inhabited the karst plateaus of the Northern Calcareous Alps corresponds to almost the entire Middle Wurmian, also called the »Alpine Cave Bear Era« (Döppes et al., 2018). ...
... Symbole: Rote Punkte und Linien bezeichnen die Mittelwerte und Fehlergrenzen der molekularen Datierung, schwarze Punkte und Linien bezeichnen die Mittelwerte und Fehlergrenzen (2σ _calBP) der Radiokohlenstoffdatierung nach der AMS-Methode. Daten nachDöppes et al., 2018;2023;Frischauf et al., 2022;Kavcik-Graumann et al., 2022, Rabeder et al., 2019 Withalm et al., 2022 und dieser Artikel (Tab. 5). ...
... The Grotta dell'Orso was also inhabited during the so-called »Alpine Cave Bear Era« -at least during the older part of this warm period. In this time period -which lasted approximately from 65,000 to 24,000 years before present (aBP) and was characterized by a warm and dry climate -almost all alpine and perialpine cave bear faunas are to be placed (Döppes et al., 2018;Nagel et al., 2018) as well as the other cave faunas in the Trieste karst plateau such as the Grotta Pocala or the Krizna jama (Calligaris et al., 2006;Döppes et al., 2018;Nagel et al., 2018;Pacher et al., 2014). ...
... The Grotta dell'Orso was also inhabited during the so-called »Alpine Cave Bear Era« -at least during the older part of this warm period. In this time period -which lasted approximately from 65,000 to 24,000 years before present (aBP) and was characterized by a warm and dry climate -almost all alpine and perialpine cave bear faunas are to be placed (Döppes et al., 2018;Nagel et al., 2018) as well as the other cave faunas in the Trieste karst plateau such as the Grotta Pocala or the Krizna jama (Calligaris et al., 2006;Döppes et al., 2018;Nagel et al., 2018;Pacher et al., 2014). ...
The fossils of the Grotta dell’Orso, recovered by Ludwig Moser (in the years 1876 to 1914), are stored in the collections of the Natural History Museum in Vienna. The cave bear remains dominating the fossil fauna were subjects to a revision. The morphological data of the dentition and the metapodials as well as the result of a DNA analysis are contradictory. The Grotta dell'Orso was possibly inhabited by two cave bear species from the Ursus spelaeus group (U. s. eremus and U. s. ladinicus) as well as by U. ingressus – similar to caves in Germany (Ach Valley) and Austria (Herdengel cave).
... Im Jahre 2001 wurden von G. Rabeder frische Knochenproben für die Datierung aufgesammelt. Die neuen Erkenntnisse über die Diversität und die ökologischen Ansprüche der alpinen Höhlenbären Bocherens et al., 2011;Döppes et al., 2018) (Ehrenberg & Sickenberg, 1929). Das Höhlenbärenmaterial umfasst fast 800 bestimmbare Reste (Tab. 1) ...
... Die Höhlenbären und Höhlenlöwen der Schreiberwandhöhle sind wichtige Klimazeugen des alpinen Pleistozäns. Die klimatischen Bedingungen, unter denen die hochalpinen Höhlenbären gelebt haben dürf-ten, wurden schon mehrfach erörtert, ebenso wie die Widersprüche zu anderen fossilführenden Sedimenten dieser Zeit (Hille & Rabeder, 1986;Döppes et al., 2011Döppes et al., , 2018Nagel et al., 2018). ...
ZUSAMMENFASSUNG Die im Jahre 1927 in der Schreiberwand-höhle (Seehöhe 2250 m) ausgegrabenen Fossilien wurden einer Revision unterzogen, um Klarheit über die taxonomische und chronologische Stellung der Höhlenbären zu erhalten. Die metrische und morphologi-sche Auswertung der Zähne und Metapo-dien spricht dafür, dass diese Fossilien zum Ladinischen Höhlenbären (Ursus spelaeus ladinicus) zu zählen sind. Von 19 Proben wiesen sechs zu geringe Kollagengehalte auf, sechs ergaben Alter jenseits von 50000 Jahren und fünf streuten zwischen rund 36000 und 49000 Jahren vor heute. Als Begleit fauna konnte der Höhlenlöwe nach-gewiesen werden sowie der Steinbock, der allerdings auch ein holozänes Element sein könnte. ABSTRACT Revision of the fossil fauna from the Schreiberwandhöhle (1543/27) in the Dachstein massif (Upper Austria) Fossils from Schreiberwandhöhle (2250 m asl.), excavated in 1927, were revised to clarify the taxonomic and chronological status of the cave bears. Metric and morphological analyses of the teeth and meta-podial bones suggest that these fossils belong to Ursus spelaeus ladinicus. Out of 19 samples taken for radiocarbon dating, six contained insufficient collagen, six yielded dates above 50000 years and the dates of the remaining five samples ranged from about 36000 to 49000 years before present. Remains of cave lion and ibex were identified as accompanying fauna; the latter, however, could also be of early Holo-cene age.
... The extremely abundant fossil record of the cave bear (Ursus spelaeus : sensu lato) from the Pleistocene of Eurasia (Kurtén 1967;Peigné et al. 2009) have provided important information on its paleobiology. Accordingly, very recently, aspects of its metabolism (Grandal d'Anglade 2018), its phylogenetic relatedness (Knapp 2018), its extinction timing (Döppes et al. 2018;Terlato et al. 2018), or even its longevity and life story (Veitschegger et al. 2018) have been successfully addressed. Despite this, the feeding preferences of the cave bear are still controversial in the literature. ...
... Our main purpose is to explore patterns of RA across maxillary teeth in the different species/subspecies of the cave bear complex and by extension their inferred feeding behaviours from this ecomorphological indicator. (Ehrenberg 1929;Abel and Kyrle 1931;Kadlec et al. 2001;Döppes and Rosendahl 2009;Döppes et al. 2011Döppes et al. , 2016Döppes et al. , 2018Pérez-Rama et al. 2011;Diedrich 2012;Horacek et al. 2012;Frischauf et al. 2014;Fortes et al. 2016;Kavcik-Graumann et al. 2016;Spötl et al. 2018;Nagel et al. 2018(in press)). As sexual dimorphism among cave bears is well reported (e.g., Kurtén 1955;Grandal d'Anglade and López-González 2005), we sexed our specimens using a protocol detailed in the Supplementary information. ...
The morphology of both crowns and tooth-roots reflects dietary specialisation in mammalian carnivores. In this article, we analyse the tooth-root morphology of maxillary teeth from CT scans of living bears (Ursus arctos, Ursus americanus, Ursus maritimus, Ursus thibetanus, Melursus ursinus, Helarctos malayanus, Tremarctos ornatus and Ailuropoda melanoleuca) in order to make inferences about the diet and feeding behaviour of the extinct cave bear (Ursus spelaeus sensu lato). Specifically, we investigate two major mitochondrial clades of extinct cave bears recognized by previous authors: Ursus ingressus and Ursus spelaeus (U. spelaeus spelaeus, U. spelaeus ladinicus, U. spelaeus eremus). Our results indicate a close association between tooth-root surface area and feeding behaviour in all living bear species. Tooth-root surface area values of cave bears suggest that they relied more on vegetative matter than living brown bears (Ursus arctos) but subtle differences between these species/subspecies could also indicate different feeding strategies among the members of cave bear complex.
... Originally, it was thought that cave bears were carnivorous due to their imposing canines. As early as 1912 O. Abel concluded from the large chewing surfaces of the molars that cave bears were herbivorous and that they used their large canines not for the acquisition of food, but as a threat and rutting weapon (Döppes et al., 2018;Frischauf et al., 2016). More recently, the herbivore character of the cave bears could be much better specified by the following methods: ...
The Hennenkopf Cave (Äußere Hennenkopfhöhle, 2,073 m a.s.l.) is located on a karst plateau named Steinernes Meer (Salzburg, Austria) which lacks a noteworthy vegetation above 1,800 m in the present time. The fossil material of the excavation from 1986 to 1991 – which is kept at the Natural History Museum in Vienna – mainly consists of isolated teeth and small bones that can be assigned to a small-sized cave bear. The chaotic stratification of the cave bear remains indicates that the cave bear remains were transported from an overlying cave section that has now disappeared. The morphological analyses of the molars show an assignment to Ursus spelaeus eremus Rabederet al., 2004. Newly analysed 14C dates now show that cave bears of the species U. s. eremus inhabited the Hennenkopf Cave in the same period as the other high-Alpine bear caves in the immediate vicinity, but also on other karst plateaus of the Northern Calcareous Alps (Nördliche Kalkalpen). Due to the extreme location of the cave in an environment that is today almost free of vegetation, these finds are impressive witnesses of a once very warm climate that allowed a lush vegetation cover at altitudes around 2,000 m above sea level.
... The samples were pretreated at roughly the same time (see Supplementary Tables S1 and S2 for details) and were measured in the same magazine in the AMS to ensure that any differences in outcome were due to the methods used rather than laboratory/instrumental background variation. Background bone samples (>50,000 BP) were pretreated and measured alongside all the samples to monitor lab-based contaminants (Döppes et al. 2019). ...
Archaeological bone collagen is highly useful for radiocarbon (¹⁴C) dating and palaeodietary reconstruction. However, collagen preservation and carbon contamination are essential considerations when extracting collagen, becoming especially crucial close to the limit of the method (50,000 years before present = BP). Strong progress has been achieved in the past two decades by ¹⁴C and stable isotopic laboratories in removing contamination from archaeological bones, but different pretreatment protocols have been proven to produce varying results. Here we compare three collagen extraction protocols used for palaeodietary studies and ¹⁴C dating, considering collagen yield, elemental and stable isotopic data, FTIR analysis, and ¹⁴C dates. We focus on the impact of ultrafiltration on the yield and quality of the extracted material. The results again underline the importance of rigorous decontamination methods to gain accurate ¹⁴C dates and demonstrate that different protocols have significant effects on the quality and yield of extracted collagen.
... In addition, there is a serious lack of reliable geochronological data of cave bear fossils before the lower limit of radiocarbon dating, i.e. older than about 45-50k cal a BP. This is unfortunate because several Alpine sites revealed bones, whose radiocarbon analyses yielded infinite ages (Pacher and Stuart, 2009;Spötl et al., 2018;Döppes et al., 2019). ...
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.
... Comparing isotopic data of cave bears with coeval Late Pleistocene large mammals, Bocherens (2018) concludes that cave bears were almost exclusively herbivorous in all analysed populations, arguing that the consumption of plants with high δ 15 N values, such as graminoids, forbs and possibly fungi, may explain the observed isotopic pattern. Döppes et al. (2018) provide new radiocarbon dates on specific populations of Alpine cave bears and accomplish a thorough revision of the 'Height Dependent Extinction Line' (HDEL), which is a compilation of the geologically most recent radiocarbon dates per altitude level. The authors conclude that a general cooling at the beginning of the Last Glacial Maximum (LGM) lead to the extinction of the cave bear in the middle of the cold stadial GS-3 (Baca et al. 2016). ...
In this issue, we cover an exceptional topic in Vertebrate Paleobiology that has been an enjoyable challenge for scientists and the popular media alike: the life and death of the Pleistocene cave bear (Ursus spelaeus). As an icon of the ice-age, the cave bear inhabited the glacial ecosystems of Eurasia, and it was the inspiration of a popular book written in 1976 by Björn Kurtén, entitled The cave bear story: life and death of a vanished animal. Although ‘The life and death’ was a summary of the knowledge acquired on cave bear biology at that time, four decades later, many aspects of its palaeoecology, extinction and evolution are still a matter of debate. With this volume, we aim to bring together the most recent research on cave bear biology in order to provide an update on the palaeoecology, biogeography, systematics, and phylogeny of this recently extinct ursine bear. We thus organised a symposium on the 1st of August 2017 as part of the three-day Annual Meeting of the European Association of Vertebrate Palaeontologists (EAVP) in Munich, Germany, that was an additional opportunity to announce the volume and to discuss this exciting subject face-to-face among specialists.
... This is a consequence of numerous studies on the morphometry (historically the most common study) on the morphodynamics of the dentition (i.e. Torres, 1988 a-f; Withalm, 2001;Rabeder, 1999Rabeder, , 2014Grandal d'Anglade and López-González, 2004;Sabol, 2005;Tsoukala et al., 2006;Baryshnikov and Puzachenko, 2011;Cvetković and Dimitrijević, 2014;Frischauf, 2014;Robu, 2016;Plichta et al., 2017), absolute dating, as well as genetics (Hofreiter et al., 2002;Orlando et al., 2002;Rabeder et al., 2004;Pacher and Stuart, 2008;Spötl et al., 2014Spötl et al., , 2017Martini et al., 2014;Döppes et al., 2016Döppes et al., , 2018Baca et al., 2016Baca et al., , 2017Fortes et al., 2017;Gretzinger et al., 2017;Terlato et al., 2018 and so on). The most important results of this knowledge have been to modify the phylogenetic tree of the cave bear. ...
Absolute dates of cave bears from Northern Italy are rare. The first radiocarbon date from Covoli di Velo Cave (Verona Province, Veneto, N. Italy) from a cave bear first phalanx is reported; its value is 29,130 ± 0.90 14C yr BP. The date, combined with morphological features of dentition suggest that cave bear populations that lived in Northern Italy were relatively underived compared to other European populations, hinting at patters of migration. Comparison of dental morphology suggest that the Covoli di Velo bear is Ursus spelaeus.
On the Hochschwab karst massif in the east of the Northern Calcareous Alps (NCA), the remains of four cave bear cubs were found in the subvertical cave Potentialschacht that opens at 2,070 m a.s.l. There is severe pathology on two individually related mandibular fragments that indicate that the cubs could have fallen into the cave through narrow gaps. There are no remains of adult bears, however. According to the morphology of the teeth, DNA analysis and 14C data, these juvenile bones and teeth belong to Ursus spelaeus eremus, which lived here at least from 46,000 to 39,000 years before present. This proves that all large plateaus of the NCA were inhabited by cave bears of Ursus spelaeus group during the so-called »Cave Bear Era« of the middle Late Pleistocene. It will be discussed, which palaeoclimatological conclusions can be drawn from it.
Juvenile cave bears (Ursus spelaeus eremus) from the Potentialschacht at Hochschwab (Styria)
Potentialschacht (1744/475) opens at an altitude of 2070 m on the northern edge of the central part of the Hochschwab plateau (Northern Calcareous Alps, Austria). The largely phreatic cave system was explored to a length of 2.3 km and a depth of 107 m. At three places, the skeletal remains of a total of four cave bear cubs, each less than one year old, were found. Serious pathologies were found on two individually contiguous mandibular fragments, suggesting that the cubs may have fallen into the cave through narrow crevices. However, there are no remains of adult bears. According to the morphology of the teeth, DNA- analysis, and 14C-data, these juvenile bones and teeth belong to Ursus spelaeus eremus, which lived here from at least 46,800 to 38,000 years before present. This shows that all major plateaus of the Northern Calcareous Alps were inhabited by cave bears of the Ursus spelaeus group during the so-called „cave bear era” of the Middle Würmian.
In the last decade, the identification of bone fragments by peptide mass fingerprinting or zooarchaeology by mass spectrometry is developing as a powerful tool in Quaternary palaeontology. The sequence of amino acids that make up the bone collagen molecule shows slight variations between taxa, which can be studied by mass spectrometry for taxonomic purposes. This requires reference databases that allow peptide identification. Although the cave bear ( Ursus spelaeus Rosenmüller, 1794) is a common component in many European Pleistocene cave sites, no peptide fingerprint taxonomic study has paid special attention to this species up to now. For peptide markers in Ursidae, the most recent proposal is based on collagen obtained from a modern brown bear sample. In this work we attempt to cover this gap by studying bone collagen of cave and brown bear samples from different origins and different chronology, applying matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry (MALDI TOF). We also performed an in-silico study of ursid bone collagen sequences published in databases. In our results we detected some discrepancies between the peptides obtained from both in silico and MALDI TOF analysis of fossil collagen and those published in the literature, in which we conclude that there are some misidentified peptides. The identification of skeletal remains by means of their peptide fingerprint is proving to be a powerful tool in palaeontology, which will bear greater fruit once the limitations of a technique that is in its initial stages have been overcome.
Abstract: On the Hochschwab karst massif in the east of the Northern Calcareous Alps (NCA), the remains of four cave bear cubs were found in the subvertical cave Potentialschacht that opens at 2,070 m a.s.l. There is severe pathology on two individually related mandibular fragments that indicate that the cubs could have fallen into the cave through narrow gaps. There are no remains of adult bears, however. According to the morphology of the teeth, DNA analysis and 14C data, these juvenile bones and teeth belong to Ursus spelaeus eremus, which lived here at least from 46,000 to 39,000 years before present. This proves that all large
plateaus of the NCA were inhabited by cave bears of Ursus spelaeus group during the so-called »Cave Bear Era« of the middle Late Pleistocene. It will be discussed, which palaeoclimatological conclusions can be drawn from it.
Weißenbach-Höhlenprojekt 108 Einleitung Braunbären hinterlassen in den Höhlen wesentlich weniger Spuren als Höhlenbären, was mit der unterschiedlichen Lebensweise zusammenhängen dürfte. Höhlenbären können als reine Pflanzenfresser die vegetationslose Zeit des Winters nur überleben, wenn sie Überwinterungsorte finden, wo sie einen energiesparenden Winterschlaf halten können: das sind die großen Höhlen, die das ganze Jahr über die gleiche Temperatur halten können, die etwa dem jährlichen Mittel-wert des Eingangsbereichs entspricht. Braunbären sind Allesfresser und können sich im Winter vor allem von Fallwild ernähren, das sie dank ihres ausgeprägten Riechsinns über große Strecken aufspüren können. Im Früh-winter halten sie eine mehrwöchige Winterruhe, "die sie auch
Zusammenfassung: Die fossile Fauna der Ochsenhalthöhle (1634/40) im Toten Gebirge-Es wird ein Überblick der bisherigen Forschungsergebnisse der fossilen Wirbeltierreste aus der Ochsenhalthöhle gegeben und die Bedeutung dieser Höhle für die Erforschung des alpinen Mittelwürms hervorgehoben. Nach der Präsentation der Basisdaten der Höhle und der wichtigsten Messdaten der Höhlenbären werden Vergleiche mit anderen Höhlenfaunen, besonders des Toten Gebirges angestellt. Das zentrale Thema dieser Studie ist die Rekonstruktion der klimatischen Bedingungen zur Zeit der Höhlenbären. Abstract: An overview of the previous research results of the fossil vertebrate remains from the Ochsenhalt Cave is given and the importance of this cave for the research of the alpine Middle Wurmian is emphasized. After the presentation of the basic data of the cave and the most important measurements of the cave bears, comparisons with other cave faunas are made, especially with caves of Totes Gebirge. The central topic of this study is the reconstruction of the climatic conditions at the time of the cave bears.
The main subject of a recent study of cave bears
from Medvedia jaskyňa Cave in the Slovenský raj
Mts. is the dentition that was found in large numbers
in the cave sediments, including hundreds of deciduous
and permanent teeth. During the field campaign
in 2007 – 2009, many isolated permanent teeth have
been excavated together with these situated still in
jaws, or in their fragments . This large number allow
representative statements about the evolutionary
level of the bears and their sex-ratios.
The variability of lower 6776 cheek teeth of spelaeoid (Ursus spelaeus, U. kanivetz, U. deningeri, U. kudarensis, U. savini and U. rossicus) and arctoid lineages (extant and extinct U. arctos) recovered from 176 palaeontological localities was analysed by methods of univariate and multivariate statistics. For comparison, teeth of U. minimus and U. etruscus species, which are ancestral taxa for cave bears and brown bear, were studied. The performed analyses indicate great variability of tooth crown in lower cheek teeth; no distinct general trend was identified in the changes, except for an increase in p4 roundness and in m3 size from U. minimus to big cave bears. Centroids of U. minimus and U. etruscus are at a great distance from those of cave bears; U. arctos occupies an intermediate position between the two groups. Molar characteristics in general are close in U. savini and U. rossicus. The bear species are differentiated in the morphospaces by morphometric variations in the lower cheek teeth but less markedly than in the morphospaces of the upper cheek teeth. A test of the inhibitory developmental cascade model showed that a linear relationship between lower molar dimensions became clear only after premolar p4 was included in the model. Evolutionary trends were found in the tooth changes with time (differently expressed in individual species and in different teeth) depending on environments, in particular on the elevation of the locality above the sea level. The allometric pattern differentiation does not always coincide with phylogenetic relationships between taxa, being most often revealed at the subspecies level. There is a dominant covariance between the upper and lower rows of teeth in accordance with the occlusion scheme. It may vary in manifestation depending on the particular species.
This study is focused on cave bears from the Grotta del Bandito (Cuneo Province, Piedmont Region, Northern Italy), especially their teeth. The morphodynamic analyses have been elaborated and compared with other cave bears from Northern Italy, and other cave bear taxa from particular European sites. The Grotta del Bandito bears are similar in size to other cave bears from Italy, and no differences were observed with the other cave bear species or subspecies (e.g. U. ingressus, U. s. ladinicus, U. s. eremus). Thus their inclusion in the U. gr. spelaeus is the most appropriate classification, and they show a simpler dental surface than the “modern” cave bears (i.e. ingressus). The question of whether a single species of spelaeus lived in North Italy, is also introduced. In fact, the differences in size are probably the result of population variability, and no clear evidence of other spelaeoid species and/or subspecies, i.e. ingressus, s. ladinicus and s. eremus, was discovered.
Ursus arctos, Ursus spelaeus eremus, bite marks on a vertebra and a fragmented os coxae of Ursus spelaeus eremus, that indicate the activity of cave lions (Panthera leo spelea) cut marks on a tibia dex.and scratch markson the surface of a radius sin., that point to human manipulation
Frischauf C., S. Krutter & G. Rabeder, Die fossile Höhlenfauna der Bärenfalle im Tennengebirge. In: Krutter S. & F. Schröder (Hrsg.), Durch die Schichten der Zeit! Neue Erkenntnisse zwischen Mesozoikum und Gegenwart. Festschrift für Erich Urbanek zum 75. Geburtstag. Forschungen des Museum Burg Golling 1 (Golling 2015), 33-44.
The fossil Fauna from the Steigelfadbalm cave was analysed by Prof. G. Rabeder and C. Rabeder MA from the University of Vienna. The cave bear bones could be attributed to the Ursus ingressus, an are thus the westernmost finds. A few chert and rock Crystal artefacts belongs to the Middle Paleolithic.
We report radiocarbon ( ¹⁴ C) dates on bone samples of Ursus ladinicus , a small cave bear species well adapted to a life in the mountains, whose remains were found in Conturines Cave. Located at 2775 m asl in the Dolomites of northern Italy, this cave is by far the highest known cave bear site worldwide. Eleven ¹⁴ C dates obtained by the Belfast and Oxford laboratories on samples showing good collagen preservation yielded consistent ages in excess of 46–50 ka BP. These results show that contrary to the previously held view these cave bear remains are older than Marine Isotope Stage 3, and likely date from a warm climate period with a high treeline, possibly the Last Interglacial.
The cave bear (Ursus spelaeus sensu lato) is a typical representative of Pleistocenemegafauna which became extinct at the end of the Last Glacial. Detailed knowledge of cave bear extinction could explain this spectacular ecological transformation. The paper provides a report on the youngest remains of the cave bear dated to 20,930 ± 140 14C years before present (BP). Ancient DNA analyses proved its affiliation to the Ursus ingressus haplotype. Using this record and 205 other dates, we determined, following eight approaches, the extinction time of this mammal at 26,100–24,300 cal. years BP. The time is only slightly earlier, i.e. 27,000–26,100 cal. years BP, when young dates without associated collagen data are excluded. The demise of cave bear falls within the coldest phase of the last glacial period, Greenland Stadial 3. This finding and the significant decrease in the cave bear records with cooling indicate that the drastic climatic changes were responsible for its extinction. Climate deterioration lowered vegetation productivity, on which the cave bear strongly depended as a strict herbivore. The distribution of the last cave bear records in Europe suggests that this animal was vanishing by fragmentation into subpopulations occupying small habitats. One of them was the Kraków-Częstochowa Upland in Poland, where we discovered the latest record of the cave bear and also two other, younger than 25,000 14C years BP. The relatively long survival of this bear in karst regions may result from suitable microclimate and continuous access to water provided by deep aquifers, indicating a refugial role of such regions in the Pleistocene for many species.
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.
The diet of the extinct European cave bear, Ursus spelaeus, has widely been debated. Diverging from the extant brown bear (Ursus arctos) approximately 1.2 million years ago, the cave bear is one of the most ubiquitous fossil bears occurring in Europe during the middle and Late Pleistocene. Early morphological studies suggested that the cave bear was likely specialized on processing tough and/or abrasive foods, while later two-dimensional low-magnification microwear studies suggested that they were omnivorous and may have consumed more bone than U. arctos. Here, we used dental microwear texture analysis (DMTA) to further interpret the diet of the cave bear. Microscopic wear features were assessed and compared to modern ursids, including the cave bears’ closest living relative, U. arctos. Results suggest that U. spelaeus consumed a diet with a diversity of textural properties, similar to most other bears and only distinguishable from the hyper-carnivorous polar bear (Ursus maritimus). Further, only U. maritimus can be distinguished from all bear species here examined (i.e., the giant panda bear, Ailuropoda melanoleuca; sun-bear, Ursus malayanus; spectacled bear, Tremarctos ornatus; American black bear, Ursus americanus; and U. arctos), with significantly greater area-scale fractal complexity (Asfc) of microwear surfaces. The DMTA of A. melanoleuca also has significantly lower Asfc than T. ornatus and U. americanus, consistent with observed dietary behavior. As modern bears vary their diets seasonally and across their range, it may be difficult to characterize the dietary ecology of extinct bears using dental microwear alone. Nevertheless, DMTA here demonstrates that U. spelaeus had a diet distinct from the hyper-carnivorous U. maritimus and instead likely consumed food with textural properties most similar to other herbivorous/omnivorous bears. Lastly, the European cave bear and North American giant short-faced bear (Arctodus simus) may have had similar diets as evident from DMTA, with U. spelaeus potentially eating tougher food items.
Count rates, representing the rate of 14 C decay, are the basic data obtained in a 14 C laboratory. The conversion of this information into an age or geochemical parameters appears a simple matter at first. However, the path between counting and suitable 14 C data reporting (table 1) causes headaches to many. Minor deflections in pathway, depending on personal interpretations, are possible and give end results that are not always useful for inter-laboratory comparisons. This discussion is an attempt to identify some of these problems and to recommend certain procedures by which reporting ambiguities can be avoided.
Tischoferhöhle and Pendling-Bärenhöhle near Kufstein, Tyrol, are among the only locations where remains of cave bear, Ursus spelaeus-group, were found in the western part of Austria. One sample from each site was radiocarbon-dated four decades ago to ca. 28 14C ka BP. Here we report that attempts to date additional samples from Pendling-Bärenhöhle have failed due to the lack of collagen, casting doubts on the validity of the original measurement. We also unsuccessfully tried to date flowstone clasts embedded in the bone-bearing sediment to provide maximum constraints on the age of this sediment. Ten cave bear bones from Tischoferhöhle showing good collagen preservation were radiocarbon-dated to 31.1–39.9 14C ka BP, again pointing towards an age underestimation by the original radiocarbon-dated sample from this site. These new dates from Tischoferhöhle are therefore currently the only reliable cave bear dates in western Austria and constrain the interval of cave occupation to 44.3–33.5 cal ka BP. We re-calibrate and re-evaluate dates of alpine cave bear samples in the context of available palaeoclimate information from the greater alpine region covering the transition into the Last Glacial Maximum, eventually leading to the demise of this megafauna.
The proposed dietary pattern of extinct Late Pleistocene cave bears (Ursus spelaeus Rosenmüller, 1794) has become controversial, as some authors have suggested that they were strictly vegetarian, whereas others maintain they were omnivores that at times ate large amounts of animal protein. We evaluated these alternatives by compiling stable isotope data of carbon (δ¹³C) and nitrogen (δ¹⁵N) from the bone collagen of adult European cave bears from the Late Pleistocene (Marine Isotopic Stage 3). The data include previously published analyses and additional data from the southeastern European (Carpathian) sites of Cioclovina, Muierii, Oase, and Urşilor. The cave bear isotopic values from bone collagen were compared with those from hair keratin occurring in grizzly bears (Ursus arctos horribilis Ord, 1815) collected from 1989 to 2009 in the western United States (Yellowstone National Park). The Yellowstone bears have access to a wide diversity of plants and animals, such that their diets can range from vegetarian to carnivorous. Thus, there was considerable δ¹³C and δ¹⁵N variation among the grizzly bear isotopic values, and the cave bear isotopic variation was encompassed within the overall grizzly bear isotopic distribution. More importantly, the δ¹⁵N distributions, reflecting principally trophic level, were not different between the cave bears and the grizzly bears; the cave bear values are, on average, slightly higher or lower than those of the grizzly bears, depending on the criteria for inclusion in the comparisons. It is therefore no longer appropriate to view Late Pleistocene cave bears as strictly or even predominantly vegetarian but as flexible omnivores within their diverse communities.
a b s t r a c t Cave bears are among the most well known extinct Pleistocene mammals. Their biogeography and taxonomy, along with the factors that led to their extinction, have been subject to long-standing con-troversy. Here, we reconstruct the phylogeography as well as the temporal and spatial population dy-namics of cave bears across their range using mitochondrial DNA control region sequences from 77 published as well as 65 new cave bear samples, Our analyses reveal a dramatic loss of genetic diversity in cave bear populations after 30,000 years before present and provide evidence for a range decline from east to west towards the onset of the last glacial maximum. Our results also suggest that the three major haplogroups within cave bears, which may correspond to distinct species, were previously more wide-spread, with relict populations in remote and alpine areas still harbouring haplotypes that have dis-appeared from most of their previous range. Applying a phylogenetic dating approach, we estimated the age of the oldest of our samples, originating from the Yana River region in north-eastern Siberia, to be around 178,000 years, which confirms a previous estimate of a Middle Pleistocene age based on its stratigraphic position. Our results extend our knowledge about the evolutionary history of cave bears,
Palaeogenetic investigations in three geographically close caves (Hohle Fels, Geißenklösterle, and Sirgenstein) in the Ach Valley near Blaubeuren (Swabian Jura) document the sudden replacement of Ursus spelaeus by Ursus ingressus around 28,000 14C BP. New radiocarbon dates suggest an earlier immigration of Ursus ingressus and at least a partial coexistence with Ursus spelaeus some 4500 years before the ultimate replacement. These two genetic types of cave bears used the same caves for hibernation and had the same herbivorous diet, as shown by the stable isotope results. In contrast, sympatric brown bears (Ursus arctos) exhibited a clearly different ecology, as shown by the carnivorous pattern of their isotopic signatures, and probably did not use the caves as dens before the Last Glacial Maximum (LGM).Once established, the younger cave bear (Ursus ingressus) remained the only cave bear for only another circa 2000 years after the last appearance of the classical cave bear (Ursus spelaeus) in the Ach Valley and elsewhere. The final appearance of cave bear (sensu lato) is now dated to 25,560 ± 130 BP, disproving a refuge area of this species in the Swabian Jura. After the extinction of cave bears (sensu lato), brown bears took over their cave dens and their nutritional niche as they shift to a diet dominated by plant food.
The European cave bear (Ursus spelaeus), which became extinct around 15,000 years ago, had several morphologically different forms. Most conspicuous of these were small Alpine cave bears found at elevations of 1,600 to 2,800 m. Whereas some paleontologists have considered these bears a distinct form, or even a distinct species, others have disputed this. By a combination of morphological and genetic methods, we have analyzed a population of small cave bears from Ramesch Cave (2,000 m altitude) and one of larger cave bears from Gamssulzen Cave (1,300 m), situated approximately 10 km apart in the Austrian Alps (Figure 1A). We find no evidence of mitochondrial gene flow between these caves during the 15,000 years when they were both occupied by cave bears, although mitochondrial DNA sequences identical to those from Gamssulzen Cave could be recovered from a site located about 200 km to the south in Croatia. We also find no evidence that the morphology of the bears in the two caves changed to become more similar over time. We suggest that the two cave bear forms may have represented two reproductively isolated subspecies or species.
The prototype mini carbon dating system (MICADAS) at ETH Zurich has been in routine operation for almost 2 yr. Because of its simple and compact layout, setting up a radiocarbon measurement is fast and the system runs very reliably over days or even weeks without retuning. The stability of the instrument is responsible for the good performance in highest-precision measurements where results of single samples can be reproduced within less than 2‰. The measurements are described and the performance of MICADAS is demonstrated on measured data.
Collagen, the organic fraction of bone, records the isotopic parameters of consumed food for carbon (δ13C) and nitrogen (δ15N). This relationship of isotopic signature between diet and tissue is an important tool for the study of dietary preferences of modern and fossil animal species. Since the first information on the isotopic signature of cave bear was reported, numerous data from Europe have become available. The goal of this work is to track the geographical variation of cave bear collagen isotopic values in Europe during Marine Isotopic Stage 3 (about 60,000–25,000 yr BP). In this study the results of new δ13C and δ15N isotopic analyses of cave bear collagen from four Central-Eastern European sites are presented, as well as a review of all published isotopic data for cave bears of the same period. The main conclusion is a lack of geographical East-West pattern in the variations of δ13C and δ15N values of cave bear collagen. Moreover, no relationship was found between cave bear taxonomy and isotopic composition. The cave bears from Central-Eastern Europe exhibit δ13C and δ15N values near the average of the range of Central, Western and Southern European cave bears. Despite the fact that most cave bear sites follow an altitudinal gradient, separate groups of sites exhibit shift in absolute values of δ13C, what disturbs an altitude-related isotopic pattern. The most distinct groups are: high Alpine sites situated over 1500 m a.s.l. – in terms of δ13C; and two Romanian sites Peştera cu Oase and Urşilor – in case of δ15N. Although the cave bear isotopic signature is driven by altitude, the altitudinal adjustment of isotopic data is not enough to explain the isotopic dissimilarity of these cave bears. The unusually high δ15N signature of mentioned Romanian sites is an isolated case in Europe. Cave bears from relatively closely situated Central-Eastern European sites and other Romanian sites are more similar to Western European than to Romanian populations in terms of isotopic composition, and probably ecology.
Isotopic tracking of carnivore palaeoecology is a relatively new approach that yielded important results for the study of the non-analogue mammoth steppe biome. After describing the prerequisite to apply this approach and the possible complications, the main achievements will be described for extinct carnivore species such as scimitar-tooth cat Homotherium serum, cave lion Panthera spelaea, giant short-faced bear Arctodus simus, cave bear Ursus spelaeus s.l., as well as for ancient representatives of extant species such as brown bear Ursus arctos and wolf Canis lupus. Isotopic tracking showed that scimitar-tooth cats in Alaska were not specialist proboscidean predators but rather generalist consumers of other large herbivores. The majority of cave lions analysed so far were focused on reindeer, some individuals were specialized on cave bears, especially in contexts of competition with cave hyenas. Giant short-faced bears in Alaska were not pure herbivores and consumed meat from reindeer, muskoxen and possibly other predators, but may have still incorporated plant resources in their menu. In contrast, all cave bear populations studied so far for which a clear dietary reconstruction could be done were virtually pure herbivores, only a few cases are still unclear. Interestingly, brown bears used the opposite extreme of the dietary spectrum when competing with other large bears such as cave bears and giant short-faced bears, i.e. were more carnivorous in Europe and more herbivorous in Alaska. Finally wolves seem to have been outcompeted by hyenas but became dominant predators during the Lateglacial in Europe to the expense of the last cave lions. The results obtained through this approach are also relevant for improving conservation strategies of endangered extant large carnivores.
The reaction conditions for the graphitization of CO2 with hydrogen were optimized for a fast production of high-quality carbon samples for accelerator mass spectrometry (AMS) measurement. The iron catalyst in use is first oxidized by heating with air to remove possible carbon and other impurities and then after evacuation reduced back to iron with hydrogen in several flushing steps to remove any iron oxide. The optimum conditions for a fast graphitization reaction were experimentally determined by changing the reaction temperatures and the H-2/CO2 ratio. The resulting graphite samples were measured by AMS to find the smallest isotopic changes (delta C-13) at a minimum of molecular fragment formation ((CH)-C-13 current). The improvements are based on thermodynamic data and are explained with Baur-Glaessner diagrams.
This study presents a cohesive review of the existing radiometric data as well as morphological and genetic analysis of bear remains from ten high-alpine caves, mostly from the Middle Würmian Interstadial complex, roughly corresponding to the marine isotope stage (MIS) 3 and dating back between 65,000–30,000 years before present. Today these caves are located in an area without any vegetation, which could not provide the herbivorous bears with sufficient food resources. It therefore can be concluded that the Middle Würmian in the Alps had to be warmer than it is today. Furthermore, congruent and conflicting data from soil formation in loess sequences as well as sinter data in caves are discussed in more detail to evaluate this hypothesis.
We present a high-resolution and independently dated multiproxylake sediment record from the paleolake at Les Échetsin southeastern France that displays synchronous changes inindependent limnic and terrestrial ecosystem proxies, in concertwith millennial-scale climate oscillations during the last glacialperiod. Distinct lake-level fluctuations, low lake organic productivity,and open, treeless vegetation indicate cold and dry conditionsin response to Heinrich events. Alternating phases of higherand low lake organic productivity, stratified surface watersand long-lasting lake ice cover, decreased or increased catchmenterosion, and tree-dominated or herb-dominated vegetation resembleDansgaard-Oeschger interstadialstadial variability. Transitionsbetween different ecological states occurred in as little as40-230 yr and seem to have been controlled by the positionof the Polar Front. Ecosystem response after 30 ka suggeststhat local climate conditions became more important. Our resultsdemonstrate that all parts of the terrestrial system respondedto the abrupt and dramatic climatic changes associated withDansgaard-Oeschger and Heinrich events, and that regional factorsmodulated ecosystem response.
Since its introduction in 19771, stable isotope analysis of bone collagen has been widely used to reconstruct aspects of prehistoric human and animal diets2–11. This method of dietary analysis is based on two well-established observations, and on an assumption that has never been tested. The first observation is that bone collagen 13C/12C and 15N/14N ratios reflect the corresponding isotope ratio of an animal's diet1–5,12. The second is that groups of foods have characteristically different 13C/12C and/or 15N/14N ratios13,14. Taken together, the two observations indicate that the isotope ratios of collagen in the bones of a living animal reflect the amounts of these groups of foods that the animal ate. Thus, it has been possible to use fresh bone collagen 13C/12C ratios to determine the relative consumption of C3 and C4 plants15–17, while 13C/12C and 15N/14N ratios have been used to distinguish between the use of marine and terrestrial foods14. The 15N/14N ratios of fresh bone collagen probably also reflect the use of leguminous and non-leguminous plants as food5, but this has not yet been demonstrated. Prehistoric consumption of these same groups of foods has been reconstructed from isotope ratios of collagen extracted from fossil bone1–11. Implicit in the application of the isotopic method to prehistoric material is the assumption that bone collagen isotope ratios have not been modified by postmortem processes. Here I present the first examination of the validity of this assumption. The results show that postmortem alteration of bone collagen isotope ratios does occur, but that it is possible to identify prehistoric bones whose collagen has not undergone such alteration.
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.
Relative warp analyses of landmarks describing cranial and mandibular shape are used for investigating patterns of morphological variation among extant bears (Mammalia, Carnivora, Ursidae) indicative of diet and feeding behavior. These patterns are used for deriving inferences about the autecology of two extinct species previously assumed to have had different dietary preferences, the North American giant, short-faced bear Arctodus simus and the Eurasian cave bear Ursus spelaeus. Results reveal a set of shared craniodental traits among the herbivorous bears, including short and vaulted skulls with well-developed zygomatic arches, lateralized orbits and small canines, concave jaws with a highly positioned condyle, large moment arms for the temporalis and masseter muscles, and long cheek teeth. In contrast, those bears that consume animal resources have long skulls with small zygomatic arches, frontalized orbits and well-developed canines, and long jaws with a deep mandibular symphysis, low muscle leverages, a condyle situated at the level of the tooth row and reduced cheek teeth. The craniodental morphology of omnivorous bears is intermediate between those of faunivores and herbivores. This is also the case of the short-faced bear and the cave bear, which suggests that previous reconstructions of the feeding ecology of these extinct species (highly carnivorous for A. simus and herbivorous for U. spelaeus) should be revised.
In the Austrian caves of Gamssulzen and Ramesch, two genetically distinct cave bears, Ursus ingressus and Ursus spelaeus eremus, apparently lived side by side for 15,000 years, together with brown bears Ursus arctos. The possible ecological partitioning of these three types of bears was investigated using multi-isotopic tracking of organic (δ13Ccoll, δ15Ncoll) and inorganic (δ13Ccarb, δ18Ocarb, δ18OPO4) fractions of bone. The cave bears from Ramesch, Ursus spelaeus eremus, were ecologically distinct from the cave bears from Gamssulzen, Ursus ingressus, both being ecologically distinct from brown bears from Ramesch, Ursus arctos. Both cave bear types were purely herbivorous but likely consumed different plant types and/or plants from different habitats, while brown bears included some animal proteins in their diet. Bone apatite δ18O values strongly suggest that both types of cave bears used isotopically distinct water sources, indicating that they may not have occupied the same landscape, either separated in space or in time due to climatic shifts. Therefore, the influence of environmental conditions strongly constrained the genetic structure of these bears.
Kiel, Universiẗat, Habil.-Schr., 1993 (Nicht für den Austausch).
Die Ochsenhalthöhle im Toten Gebirge (Steiermark)
- C Frischauf
Frischauf C 2011. Die Ochsenhalthöhle im Toten Gebirge (Steiermark).
Unpub. Master thesis. Univ. Vienna.
New scientific results from the Arzberg Cave near Wildalpen
- D Döppes
- M Pacher
- C Frischauf
- G Rabeder
Döppes D, Pacher M, Frischauf C, Rabeder G. 2012. New scientific
results from the Arzberg Cave near Wildalpen, Styria, Austria.
Braunschw Naturk Schr. 11:41-48.
Golling: Forschungen des Museums Burg Golling
- Geburtstag
Geburtstag). Golling: Forschungen des Museums Burg Golling; p.
33-44.
The cave bear fauna (Ursidae, Mammalia) from Steigelfadbalm near Vitznau
- C Frischauf
- E Nielson
- G Rabeder
Frischauf C, Nielson E, Rabeder G. 2018. The cave bear fauna (Ursidae,
Mammalia) from Steigelfadbalm near Vitznau. Canton of Lucerne
(Switzerland): Acta Zoologica Cracoviensia (in press).
Die Gamssulzenhöhle im Toten Gebirge
- G Rabeder
Rabeder G. 1995. Die Gamssulzenhöhle im Toten Gebirge. Mitt Komm
Quartärforsch Österr Akad Wiss. 9:1-133.
The cave bears from Schlenken-Durchgangshöhle (Schlenken Passage Cave Osterhorn Massif
- T Knaus
- B Schopf
- C Frischauf
- N Kavcik-Graumann
- G Rabeder
Knaus T, Schopf B, Frischauf C, Kavcik-Graumann N, Rabeder G. 2017.
The cave bears from Schlenken-Durchgangshöhle (Schlenken Passage
Cave Osterhorn Massif, Salzburg, Austria). Aragonit. 22(1):13-14.
Ursus spelaeus from Grotta supra Fontana Marella
- R Perego
- E Zanaldi
- A Tintori
Perego R, Zanaldi E, Tintori A. 2001. Ursus spelaeus from Grotta supra
Fontana Marella, Campo di Fiori Massiv (Varese, Italy): morphometry and Palaeoecology. Rev. Ital. Pal. 107:451-462.
Cave bear site Jama pod Herkovina pecmi (N. Slovenia): preliminary study
- V Pohar
- I Debeljak
- G Rabeder
Pohar V, Debeljak I, Rabeder G 2003. Cave bear site Jama pod
Herkovina pecmi (N. Slovenia): preliminary study. 9ème
Die Höhlenbären von Conturines. Entdeckung und Erforschung einer Dolomiten-Höhle in 2800 m Höhe
- G Rabeder
Rabeder G. 1991. Die Höhlenbären von Conturines. Entdeckung und
Erforschung einer Dolomiten-Höhle in 2800 m Höhe. Bozen: Athesia.
Radiocarbon Constraints on the Age of the World's Highest-Elevation Cave-Bear Population
- C Spötl
- P Reimer
- G Rabeder
- R C Bronk
Spötl C, Reimer P, Rabeder G, Bronk RC. 2017. Radiocarbon Constraints
on the Age of the World's Highest-Elevation Cave-Bear Population,
Conturines Cave (Dolomites, Northern Italy). Radiocarbon. 1-9.
doi:10.1017/RDC.2017.60
Chronological and Isotopic data support a revision for the timeing of cave bear extinction in Mediterranean Europe
- G Terlato
- H Bocherens
- M Romandini
- N Nannini
- H Ka
- M Peresani
Terlato G, Bocherens H, Romandini M, Nannini N, Ka H, Peresani M.
2018. Chronological and Isotopic data support a revision for the
timeing of cave bear extinction in Mediterranean Europe. Hist Biol.
1-11. doi:10.1080/08912963.2018
Die Ramesch-Knochenhöhle im Toten Gebirge. Mitt Komm Quartärforsch Österr Akad Wiss
- P Hille
- G Rabeder
Hille P, Rabeder G. 1986. Die Ramesch-Knochenhöhle im Toten
Gebirge. Mitt Komm Quartärforsch Österr Akad Wiss. 6:1-66.