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The cave bear fauna of the cave Schottloch (Dachstein Mountains, Austria)

Authors:
Nadja Kavcik-Graumann at University of Vienna
Nadja Kavcik-Graumann
Federica Alberti at Universität Potsdam and Reiss Engelhorn Museum Mannheim
Federica Alberti
  • Universität Potsdam and Reiss Engelhorn Museum Mannheim
Doris Döppes
Doris Döppes
Doris Döppes
  • This person is not on ResearchGate, or hasn't claimed this research yet.
Ronny Friedrich at Curt Engelhorn Zentrum Archäometrie
Ronny Friedrich
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Abstract and Figures

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.
Ground plan and longitudinal section of the bear cave »Schottloch« at the southern edge of the plateau »Auf dem Stein« after Döppes & Rabeder, 1997: 225. | Abb. 1: Grundriss und Längsschnitt der Bärenhöhle »Schottloch« am Südrand des Plateaus »Auf dem Stein« nach Döppes & Rabeder, 1997: 225.
Ground plan and longitudinal section of the bear cave »Schottloch« at the southern edge of the plateau »Auf dem Stein« after Döppes & Rabeder, 1997: 225. | Abb. 1: Grundriss und Längsschnitt der Bärenhöhle »Schottloch« am Südrand des Plateaus »Auf dem Stein« nach Döppes & Rabeder, 1997: 225.
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Geographic location of the karst plateaus »Auf dem Stein« and »Gosauer Stein« in the Dachstein Mountains, with position of the bear caves Schreiberwand cave (Sr) and Schottloch (SchL). Map sketch by G. Rabeder and N. KavcikGraumann. | Abb. 2: Geographische Lage der Karstplateaus »Auf dem Stein« und »Gosauer Stein« im Dachsteingebirge mit der Lage der Bärenhöhlen Schreiberwandhöhle (Sr) und Schottloch (SchL). Kartenskizze von G. Rabeder und N.
Geographic location of the karst plateaus »Auf dem Stein« and »Gosauer Stein« in the Dachstein Mountains, with position of the bear caves Schreiberwand cave (Sr) and Schottloch (SchL). Map sketch by G. Rabeder and N. KavcikGraumann. | Abb. 2: Geographische Lage der Karstplateaus »Auf dem Stein« und »Gosauer Stein« im Dachsteingebirge mit der Lage der Bärenhöhlen Schreiberwandhöhle (Sr) und Schottloch (SchL). Kartenskizze von G. Rabeder und N.
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Morphological index of the fourth premolars (p4/4 index) of alpine cave bear faunas in correlation to the altitude of the cave entrances. Abbreviations: BF = Bären-falle, RK = Ramesch-Knochenhöhle, SchL = Schottloch, Sr = Schreiberwandhöhle. | Abb. 5: Morphologischer Index der vierten Prämolaren (p4/4-Index) alpiner Höhlen-bärenfaunen in Korrelation mit der Höhenlage der Höhleneingänge. Abkürzungen: BF = Bärenfalle, RK = Ramesch-Knochenhöhle, SchL = Schottloch, Sr = Schreiberwandhöhle.
Morphological index of the fourth premolars (p4/4 index) of alpine cave bear faunas in correlation to the altitude of the cave entrances. Abbreviations: BF = Bären-falle, RK = Ramesch-Knochenhöhle, SchL = Schottloch, Sr = Schreiberwandhöhle. | Abb. 5: Morphologischer Index der vierten Prämolaren (p4/4-Index) alpiner Höhlen-bärenfaunen in Korrelation mit der Höhenlage der Höhleneingänge. Abkürzungen: BF = Bärenfalle, RK = Ramesch-Knochenhöhle, SchL = Schottloch, Sr = Schreiberwandhöhle.
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Chronological data of cave bear remains (»plateau bears«, Ursus spelaeus) from the large bear caves of the Northern Calcareous Alps in relation to the altitude of the caves. Key to symbols: red dots and lines denote the mean values and error limits of molecular dating, black dots and lines denote the mean values and error limits (2σ _calBP) of radiocarbon dating by the AMS method. Data after Döppes et al., 2018; 2023; Frischauf et al., 2022; Kavcik-Graumann et al., 2022, Rabeder et al., 2019; Withalm et al., 2022 and this volume (Tab. 5). | Abb. 6: Chronologische Daten von Höhlenbärenresten (»Plateaubären«, Ursus spelaeus) aus den großen Bärenhöhlen der Nördlichen Kalkalpen in Abhängigkeit von der Höhenlage der Höhlen. 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 nach Döppes 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).
Chronological data of cave bear remains (»plateau bears«, Ursus spelaeus) from the large bear caves of the Northern Calcareous Alps in relation to the altitude of the caves. Key to symbols: red dots and lines denote the mean values and error limits of molecular dating, black dots and lines denote the mean values and error limits (2σ _calBP) of radiocarbon dating by the AMS method. Data after Döppes et al., 2018; 2023; Frischauf et al., 2022; Kavcik-Graumann et al., 2022, Rabeder et al., 2019; Withalm et al., 2022 and this volume (Tab. 5). | Abb. 6: Chronologische Daten von Höhlenbärenresten (»Plateaubären«, Ursus spelaeus) aus den großen Bärenhöhlen der Nördlichen Kalkalpen in Abhängigkeit von der Höhenlage der Höhlen. 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 nach Döppes 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).
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The karst plateau »Auf dem Stein« in the eastern Dachstein Mountains, view from the Gjaidalm to SE, is today overgrown with elements of the subalpine and alpine vegetation stages. The red arrow marks the location of the bear cave Schottloch. | Abb. 7: Das Karstplateau »Auf dem Stein« im östlichen Dachsteingebirge mit Blick von der Gjaidalm nach Südosten, auf dem heute Pflanzen der subalpinen und alpinen Vegetationsstufe wachsen. Der rote Pfeil markiert die Lage der Höhle Schottloch.
The karst plateau »Auf dem Stein« in the eastern Dachstein Mountains, view from the Gjaidalm to SE, is today overgrown with elements of the subalpine and alpine vegetation stages. The red arrow marks the location of the bear cave Schottloch. | Abb. 7: Das Karstplateau »Auf dem Stein« im östlichen Dachsteingebirge mit Blick von der Gjaidalm nach Südosten, auf dem heute Pflanzen der subalpinen und alpinen Vegetationsstufe wachsen. Der rote Pfeil markiert die Lage der Höhle Schottloch.
… 
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of Museum Burg Golling

Nadja Kavcik-GraumannFederica AlbertiDoris DöppesRonny FriedrichJulia Stockhammer
Susanne LindauerMichael HofreiterGernot Rabeder
-
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--
--

The cave bear fauna of the cave Schottloch
(Dachstein Mountains, Austria)
Ursus spelaeus eremus 
Ursus spelaeus eremus 
Abstract:            
               
 
    
             
Ursus
spelaeus eremus     
       
            
             
 

Kurzfassung           

      
           
       
  
      Ursus spelaeus eremus
           
         

           
      


-
Introduction
        
        
   
         
-
        
         
  
        
        
     
        
         
    -
           
Fig. 1-2

          
  

         Fig. 3 
       
         
Canis lupus

         

    
          
        
-    
  
          
  
          

     
          
       -

Basics



Fig. 1             
          
Abb. 1         

Fig. 2   
       
      
     
    -
  Abb. 2   
     
      

   
-

-
Material

 
        
      

  
   
      
      
        

         

       
         
     

Methods


         
      
       
    
    
Fig. 4 
       

    
        
-
-     -
       
      

Results
Metric of canines, sex index

           
    
 Tab. 1     Fig. 3  

Metric of cheek teeth and metapodia

         
Ursus ingressus      Tab. 2
         
       
Tab. 3
Fig. 3       
  Abb. 3- 
-
Fig. 4         
        
            
    Abb. 4-    
   -    


Tab. 1            
Tab. 1
 -     

Canini crown length crown width number length stand. with stand.
     
     
     

    
 

-
LHD-diagram
-


-
         
        
    Ursus ingressus  

Ursus spelaeus  U. spelae-
us eremusU. spelaeus ladinicus 
Fig. 4

Ursus spelaeus        

        
  

         
       

Morphology of premolars
Tab. 4 
          


         
      

     Ursus spelaeus eremus Fig. 5

          
U. ingressus U. spelaeus
Fig. 5        
 
   -       
Abb. 5      -  

-
Element length width md index

   
   
  

   
   
  


  
  
 

  
  
 

   
   
 

   
   
  

  
  
 
-   
 
   
 
Tab. 2 Ursus s. eremus
           
 Tab. 2
     Ursus s. eremus    
          

Element n length dew PI length
stand.
dew
stand. PI stand.
       
       
       
       
       
      
      
       
       
       

    
Tab. 3U rsus s. eremus
              
               
 Tab. 3    Ur sus s.
eremus          



-
Chronology of Schottloch bears

          
    
          
         
         
 
          
Tab. 5
Discussion

  
  
    
          

        
Ursus spelaeus  U.
spelaeus eremus        
      Ursus s.
ladinicus
      
       
         
   -
  
          
   
      Ursus
ingressus         

 


   -      
         

          
    U. spelaeus     
U. ingressus 
       

     

Tab. 5   
  Fig. 6    





The vegetation on the karst plateaus in the
alpine cave bear period
        
   
Fig. 6         Ursus spelaeus


    σ
             
     -       
         Abb. 6   
Ursus spelaeus 
     
         
 
σ _     -  
         -   

Morphotypes A A/B A/D B1/C1 B2 C1 C1/C2 C2 C3 D1 D1/2 D2 n

 
Morphotypes B1/C1 B2 C1 C1/C2 C2 C3 D1 D1/2 D2 n
 
Morphotypes A A/B A/D n

Tab. 4 Ursus s. eremus  
Tab. 4Ursus s. eremus 

-
MAMS sample name bone element 14C age error ± δ13C 2σ calBP 2σ calBP mean C:N C (%) collagene (%) error ±
 -    - -     
 -    - -     
 -    - -     
 -    - -     
 -    - -     
 -    - -     
 -    - -     
 -    - -     
 -      
Tab. 5  Ursus s. eremus   Tab. 5 Ursus s. eremus

   

           
    

         
    
         
 -

            
         

        
          
           
         
        - 

Conclusion
          
   Ursus spelaeus     -
 Fig. 4      Fig. 5  
U. spelaeus eremus U. s. ladinicus  
   
        
         
    -  
Fig. 5      Ursus spelaeus eremus
        
U. s. eremus
Taxonomy, chronology and diet of the so called »plateau bears«

         
      


            -
Ursus spelaeus eremus Ursus spelae-
us ladinicus Ursus ingressus 
   Fig. 4-5     

         
     U. ingressus  U. s. eremus
          
         
-
Fig. 7 
      Abb. 7    


-
        
          
          
        

           

   -       

Acknowledgments

   
 -        

  

References
        
      
 
-
      - 
        
-
     -

        -
   

      
 
 
    
-

 
-
-    
        
        
   -   
-   
     
-
   -   
         

 -
-       
        
 -
-      


 


Ursus spelaeus  -
      
-
-
  --
     
       


--

      -
-  

      
-
     
-
    
 --

       
--
        
Ursus spelaeus 
- --
    -   
       -
    
--
     
     


      Ursus spelaeus
 
-

- -

  -
Ursus
spelaeus eremus
 --
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Juvenile Höhlenbären (Ursus spelaeus eremus) aus dem Potentialschacht am Hochschwab (Steiermark)
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  • May 2022
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.
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The cave bear fauna of Schottloch (Dachstein mountains, Austria) and the cave bears of the great karst plateaus of the Northern Alps
Poster
Full-text available
  • Sep 2022
The study of the fossil fauna from the Schottloch in the Dachstein Mountains is taken as an opportunity to give an overview of all cave bear faunas described and dated so far in the Northern Alps. From the metrical and morphological data of the cave bear remains it can be concluded that on the one hand the cave bears of the Schottloch taxonomically correspond to the other bears of the large karst plateaus of the Northern Limestone Alps and on the other hand there are major differences to Ursus ingressus, which dominates especially in the lower situated bear caves. The comparison of the altitudes of all important cave bear sites gives a surprisingly unanimous picture: the caves with Ursus spelaeus eremus and U. s. ladinicus are located at altitudes between about 1600 and 2350 m.a.s.l., while all bear populations with U. ingressus are below 1400 m.a.s.l. Thus, the two cave bear groups differ not only genetically but probably also in their adaptation to mountain life. However, they have lived simultaneously in the same mountains for a period of more than 30,000 years.
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Weißenbach-Höhlenprojekt The Fossil Fauna of the Ochsenhalthöhle (1634/40) in the Totes Gebirge
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Full-text available
  • Apr 2022
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.
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Functional morphology of the cave bear (Ursus spelaeus) mandible: a 3D geometric morphometric analysis
Article
Full-text available
  • Oct 2015
The diet of the fossil cave bears (Ursus spelaeus group) has been debated extensively. Thought traditionally to be herbivorous, more recent studies have proposed more meat in the cave bear diet. To test this, the mandibular morphology of cave bears was analysed using 3D geometric morphometrics and compared to that of extant Ursidae. Landmarks for 3D digitisation of the mandible were chosen to reflect functional morphology relating to the temporalis and masseter muscles. Extant and extinct Pleistocene Ursidae were digitised with a MicroScribe G2. Generalised Procrustes superimposition was performed, and data were allometrically and phylogenetically corrected. Principal component analysis (PCA), two-block partial least squares analysis (2B-PLS), regression analysis and discriminant function analysis were performed. PCA and 2B-PLS differentiate between known dietary niches in extant Ursidae. The lineage of the cave bear runs parallel to that of the panda (Ailuropoda melanoleuca) in morphospace, implying the development of morphological adaptations for eating foliage. A regression of shape onto foliage content in the diet and a discriminant function analysis also indicate that the cave bear diet consisted primarily of foliage.
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Morphological responses of cave bears (Ursus spelaeus group) to high-alpine habitats
Article
Full-text available
  • Jan 2008
A comparison of metrical and morphological data from more than 30 bear populations belonging to the cave bear group (U. spelaeus and its relatives U. eremus, U. ladinicus, U. ingressus as well as their predecessor U. deningeri) shows that the different species developed very different adaptations to the altitude of their habitats. Whereas there is a reduction of body size in Ursus eremus and U. ladinicus correlated with the altitude of the habitat (“alpine nanism”) no such correlation can be observed in U. ingressus. In U. ingressus there is a positive correlation of the tooth indices and the altitude of the habitat, i.e. in the higher sites the teeth are more evolved.
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Evidence for Reproductive Isolation between Cave Bear Populations
Article
Full-text available
  • Feb 2004
  • CURR BIOL
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.
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Extinction pattern of Alpine cave bears - new data and climatological interpretation
Article
  • Jun 2018
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.
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Pliozäne und pleistozäne Faunen Österreichs
  • Jan 1997
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  • D Döppes
  • G Rabeder
Döppes, D. & Rabeder, G. (1997): Pliozäne und pleistozäne Faunen Österreichs. Ein Katalog der wichtigsten Fossilfundstellen und ihrer Faunen. -Mitt. Komm. Quartärforsch. Österr. Akad. Wiss. 10: 1-411.
  • Jan 1962
  • 394-437
  • K Ehrenberg
Ehrenberg, K. (1962): Bemerkungen über die Bestände an Höhlenfunden im Oberösterreichischen Landesmuseum. -Jahrbuch Oberösterreichischer Musealverein 107: 394-437.
Atlas of Cave Bear Osteology
  • Jan 2021
  • 1-138
  • C Frischauf
  • G Rabeder
Frischauf, C. & Rabeder, G. (2021): Atlas of Cave Bear Osteology. -Naturkundliche Mitteilungen aus den Landessammlungen Niederösterreich 30: 1-138.