It was possible to study the material with the support of the City of Balve as the owner, and Dr. Schumacher-Mathäus. Comparative material was studied in different collections. I thank Mag. J. Törnig-Struck from the Museum für Stadt- und Kulturgeschichte Menden for the possibility to study the Keppler Cave material, and Dr. W. Bleicher for support. Free access to the collection of the Museum für Naturkunde Berlin of Humboldt University was possible due to the kindness of PD Dr. O. Hampe. Lion material studies in the Goldfuss-Museum of the Rheinische Wilhelms-University Bonn were supported by Dr. G. Heumann. The curator of the Pleistocene collection in the British Museum of Natural History London, A. Currant, gave access to Pleistocene lion material, especially the skull from the German Perick Caves. Dr. M. Bertling from the Geologisch-Paläontologische Museum der Westfälischen Wilhelms-Universität Münster made the study of many finds of the Sauerland and Münster Bay possible, which are housed in the Geological-Palaeontological Museum’s collection of the Westphalian Wilhelms-University Münster. The museum’s head of the Naturkundemuseum Siegsdorf, Dr. R. Darga, allowed the comparison of the Upper Pleistocene lion skeleton from the open air mammoth site Siegsdorf (Bavaria, South Germany). The museum’s head of the Staatliche Naturhistorische Sammlungen Dresden, Dr. U. Linnemann, supported the study of the collection from the Perick Caves. Finally, bone material from the hyena open air prey depot site Herten-Stuckenbusch and a modern African lion skeleton in the collection of the Museum for Ur- und Ortsgeschichte Quadrat Bottrop could be compared with the kind support of M. Walders. A thankful first critical review was made by Dr. H. O’Reagan. Finally, I thank the reviewers Dr. P. Auguste and Dr. A. Argant for the critical reviews and supporting comments. The company PaleoLogic sponsored the work, which was illustrated by a lion picture of G. “Rinaldino” Teichmann.
1Historically collected in caves, late Pleistocene bones, which were not often identified as lion bones, have been rediscovered from the Sauerland Karst cave and bone-rich region of north-western Germany, including the here focussed Balve Cave. These bones were found during many pioneer cave excavations during the 19th century (Virchow, 1870; von Dücker, 1870; von Dechen, 1871, 1879; Kremer, 1873; Schaafhausen, 1875; Zygowski, 1988; Ahrweiler, 1995; fig. 1A-B).
2The horizontal and large, but only 90 m long Balve Cave (elevation 230 m, fig. 1C) splits after 54 m into two branches: the Virchow and the Dechen branches, named after their excavators (fig. 1D). With its 11 m high and 18 m wide portal (fig. 1E), it is easily visible from the Hönnetal (Hönne Valley). At the beginning of the 19th century, the Balve Cave began to be intensively excavated by farmers for bones to use those as phosphatic agents. During that time, the uppermost speleothem layer, which is still today attached 4-5 m above the ground level to the cave walls, and the upper 1-2 meters of sediment of the upper “culture layers” were completely removed. First excavations concerning archaeology and faunal remains were made in 1843/44 (Bahnschulte, 1940). After this, Virchow (1870) and von Dechen (1871) excavated and described new finds from different areas of the cave. Although Kramer (von Dechen, 1879) and von Dücker (1870) also excavated in the cave, the largest historical scientific explorations were made by Bahnschulte (1940). He worked with the use of systematic trenches, which have produced the most important stratigraphic information (fig. 2). Finally, Günther (1964) excavated in 1959 close to the old trenches of Bahnschulte.
3The Balve Cave is now one of the most famous caves in the Sauerland Karst of northern Germany (Brandt, 1939), but also a well-known archaeological locality in Europe. It has a combination of animal and human use over a period of more than 120,000 years and contains both, a large amount of artefacts (about 53,000), and bones (about 2,000) from several layers (Günther, 1988). The cave became famous early, because of its rich and large amount of Lower and Middle Palaeolithic (Acheulean, Mousterian, Micoquian) lithic artefacts, mainly made of regional black lydite and few imported quartzite or nordic flintstone (Kremer, 1873; Bahnschulte, 1938, 1940; Beck, 1975), as well as several bone artefacts that were found and repeatedly studied (Andrée, 1924, 1928; Brandt, 1939; Günther, 1964, 1988; Bosinski, 1984; Järis, 1992, 1993). A horse engraving from the Aurignacian period is the only important art find (Andrée, 1930).
4Micromammals were already sieved in the 19th century and listed by Farwick (1870), von der Marck (1872) and Nehring (1879a, 1879b, 1880, 1883), but a modern stratigraphy-related and palaeoclimatologic micromammal analysis is missing. The macrofauna was generally only mentioned or listed without a good stratigraphic context, detailed species or subspecies names or quantitative analyses (Virchow, 1870, 1879; von Dücker, 1870; von Dechen, 1871, 1879, 1884; Kremer, 1873; Wollemann, 1887). At least Pohlig (1888) described woolly mammoth Mammuthus primigenius remains, but also not in detail, and not with a taphonomic study about hyena bite marks on those bones, or with a discussion to its possible hyena prey origin, which was later proven for some chewed and bite damaged bones (Diedrich, 2011d). A rare musk ox Ovibos moschatus skull (Andrée, 1933) was reported, and is now preserved in the small regional Balve Museum. Hippopotamus amphibius was described only briefly (Wollemann, 1887), which indicates the presence of interglacial (Eemian) sediments and fauna. The Pleistocene macrofauna was generally not studied and only listed by non-palaeontologists (Bahnschulte, 1940; Günther, 1988). This also results of the artefact and bone spreading in many collections (see Material and methods) and of problematic stratigraphic records. Hence, it was generally thought all to be “human bone importation” (Andrée, 1928; Günther, 1964, 1988). Only cave bear remains were mentioned as being the non-imported faunal remains of a cave bear den site (Bahnschulte, 1940).
513 cave sites in the Sauerland Karst and five open air sites are known in Westphalia to have provided Upper Pleistocene Panthera leo spelaea (Goldfuss 1810) bone material (fig. 1A-B; Diedrich, 2009d) from Perick Caves hyena and cave bear den including the only preserved complete skull of the Sauerland Karst caves (60 bones; Diedrich, 2009a), Bilstein Caves cave bear den (30 bones; Diedrich, 2009d). The Balve Cave is also one of the few north-western German caves with a higher amount of lion remains as described herein. In this study, 48 lion bones from Balve Cave, and 35 bones from Keppler Cave cave bear den are added in a review, such as a few more remains from the Grürmanns Cave hyena den, Johannes Cave hyena den, Kreuz Cave, Martins Cave cave bear and hyena den, Teufelskammer Cave hyena den (Diedrich, 2011b), and Wilhelms Cave hyena den (fig. 1B). This lion material described herein and its taphonomic context in hyena dens and cave bear dens is of importance for the palaeobiological understanding of late Pleistocene lions and their antagonism to the last Ice Age spotted hyenas of Europe, cave bear hunt by predators (hyenas/lions; Diedrich, in press), and the taphonomy of lion bones in caves in general (cf. Diedrich, 2009a, 2009b). The material especially from the Balve Cave will be discussed in its “non-human context”, whereas the former discussion about “importation of carnivores” by human hunters to their camp sites will be critically discussed with the material from a “human cave site” and “non-human cave sites” of the Sauerland caves.
Fig. 1 A : Cave and open air sites with Upper Pleistocene lion material.
The most important are the Bottrop open air site along with the Perick Caves, Wilhelms Cave, Keppler Cave, Bilstein Caves and Balve Cave. Other localities contain only a single or few Panthera leo spelaea bone remains.
6
Fig. 1 B : Extension of compared areas all over Germany and Czech Republic.
7
Fig. 1 C : Balve cave portal from historical times (post card).
8
Fig. 1 D. Balve cave map with the Dechen and Virchow branches.
Fig. 1 E. Modern photo of the Balve cave entrance portal.
Fig. 2: Generalized stratigraphy of the seven meters thick section of the balve cave (NW Germany) and its megafauna content.
Stratigraphy reconstructed after Virchow (1870), von Dechen (1871), Andrée (1928), Bahnschulte (1940) and Günther (1988)
9The studied Sauerland Caves are all dated from the early to middle Upper Pleistocene by the megafauna assemblages (hyena bone accumulations, cave bear den bone accumulations), and cave bear tooth morphologies at the Perick, Martins, Grürmanns, Balve, Bilstein, and Keppler Caves (method after Rabeder et al., 2000), whereas Pleistocene radiocarbon data are still missing.
10The Balve Cave was mostly historically excavated, and exact stratigraphic documentation, especially of the here-described bone material, is then missing. At least a coarse stratigraphy of a seven meter thick section composed after the descriptions of Virchow (1870), von Dechen (1871), Andrée (1928), Bahnschulte (1940) and Günther (1988) is reproduced here with some new faunal, climatic and cave occupation interpretations (fig. 2). Whereas Bahnschulte (1940) used the Horizon numbers I to V, Günther (1964, 1988) differentiated those former divided I-III layers into the layers 1 to 6 (fig. 2).
11The oldest layer in the cave is a basal clay which might even be deposited from pre-Pleistocene time. No faunal remains were observed.
12The loams of the first Middle Pleistocene layer that contain here-determined early forms of cave bears with Ursus deningeri (material in the Museum of Humboldt University in Berlin) is of possible late Middle Pleistocene age. The cave was used by early cave bears as a cave bear den cave. At this time, no human impact can be verified. Therefore, this oldest bone layer might even be older than Saalian, and is most likely even of pre-Saalian age. The following four similar layers consisting of non-rounded limestone gravels are dated as Saalian. A Middle Pleistocene age can be estimated, but again, these layers might even be older. At least, they represent a cold period stage. From those layers, cave bears are present, most probably still with U. deningeri forms, for which the few materials cannot yet be analysed more in detail. These bone layers also contained Canis, and possibly Praemegaloceros as well. Even a medium-sized Panthera pardus felid can be added to the fauna with a single canine tooth. Perhaps, this cave was at that time also already used by hyenas as a den site, to which imported prey remains of the older layers might be attributed.
13The first layer with artefacts is the Balve I Horizon, which is separated by layers 5 and 6. The older layer 6 (30-50 cm) is a dark-brown sandy clay containing a few small, and some large, weathered, and rounded limestone pebbles. This layer, which contains late Acheulean artefacts, is of Eemian interglacial age, which is also supported by pollen analyses (Günther, 1988). It was most likely from these layers that Hippopotamus amphibius was found (Wollemann, 1887).
14The younger layer 5 (20-50 cm) is a red-brown sandy clay with some non-rounded limestone pieces. From these layers, cave bears (Ursus cf. spelaeus subsp.) and Coelodonta antiquitatis are reported (Bahnschulte, 1940). These woolly rhinoceros bones have been redescribed to be mostly of hyena den bone accumulation origin (Diedrich, 2011a). The age of this layer seems to be at the beginning of the early Weichselian.
15The Balve II Horizon is subdivided into layers 3 and 4. Layer 4 is grey-yellow clay with sharp-edged 1-20 cm large limestones. The rich megafauna indicates a cold climate: Mammuthus primigenius, Bison priscus, Equus caballus cf. przewalskii, Megaloceros giganteus, Crocuta crocuta spelaea, Ursus cf. spelaeus subsp. and Canis lupus cf. spelaeus. Several cranial and postcranial bones of the here-described Panthera leo spelaea material must originate from those layers. Layer 3 is small grained limestone gravel (6-12 cm in size), which seems to represent the first maximum cold peak of the early Weichselian. Even larger smashed bones from mammoth, which were used as burning material (“bone coal”) instead of wood, are present in Horizon I that contains a higher amount of artefacts from the early Micoquian (Günther, 1988).
16The Balve III Horizon was subdivided into a-c. The oldest, IIIa Horizon, is similar to layers 1 and 2. Layer 2 (20-40 cm thick) is made of clay with a few stones. Above it, layer 1 (1-5 cm thick) is represented by light to dark, grey soil, in which small, to coarse, non-rounded limestone pieces are present. Here “bone coal” and artefacts (late Micoquian) are present. Especially from this layer, all three carnivores Ursus cf. spelaeus subsp., Crocuta crocuta spelaea (cf. Diedrich, 2011a) and Panthera leo spelaea are reported historically, as well as the typical cold megafauna from the early Weichselian, such as M. primigenius, C. antiquitatis, E. caballus cf. przewalskii, E. hydruntinus, M. giganteus and R. tarandus. Even some chew marks from the porcupine Hystrix (A.) brachyura can be included here. They are similar to the rodent gnaw mark shapes recently described for chewed bones from the German open air hyena den site Rote Berg Saalfeld and Czech caves of overlapping European hyena and porcupine dens (Diedrich, 2009c). This layer can be incorporated here to represent hyena activity, den use, and prey importation, because some bones with chew marks and incomplete woolly rhinoceros bones can be referred to this layer (Diedrich, 2011a). The IIIb Horizon (30-50 cm thickness) is made of brown-yellow clay with some rounded limestone pebbles, many artefacts (late Micoquian), and animal bones. In the IIIc Horizon (80-100 cm) the grey sediments another culture layer is present. This stratum seems to be of middle Weichselian age (Günther, 1988).
17A brown-yellow loam with a few stones (60-100 cm thick) was called Balve-Horizon IV. It is interrupted in the middle by a 30 cm thick, grey and coal-rich culture horizon. This level is fauna- and artefact-rich. A late Mousterian, and on the top a “Moustérien à lames” lithic inventory has been described, dating into the middle Weichselian (Günther, 1988).
18The last preserved Balve Horizon V was described as light yellow-grey clay that contained a few small stones. Here, artefacts from the modern human periods of the Aurignacian were figured (Bahnschulte, 1940; Günther, 1988). Therefore, the boundary between the middle and late Weichselian can be set. In these layers, cave bear remains were also excavated, plus a still-mixed mammoth steppe and intermediate fauna. From this Aurignacian period, a horse head engraving was published as the only art find (Andrée, 1933).
19Younger modern human archaeological layers of a Magdalenian period were destroyed in the past and sediments or sections are missing from the “Reindeer horizon” (cf. Günther, 1988). As compared to historical descriptions and still present bone material, the typical megafauna featuring the three large carnivores Panthera, Ursus and Crocuta is absent. In those final Weichselian layers, there is an abundance of reindeer Rangifer tarandus remains, including many dropped antlers that have no carnivore bite or chew marks. Those must have been collected and accumulated by humans and not by hyenas or other carnivores (wolves). Additionally Equus caballus cf. przewalskii, horses (see classification in Zessin et al., 2009) and wolves are present, as is finally the only musk ox Ovibos moschatus remain (Andrée, 1933), which fits to the maximum cold period, and periods after this, of the late Weichselian. Artefacts seem to be all of the final Magdalenian time of the V and VI stages (Günther, 1988).
20The late Palaeolithic horizon was covered by a thick speleothem layer, which was dated between 8,000-10,000 yr BP (Zygowski, 1988), but for which a new modern dating is still lacking.
21Above this layer Neolithic and early pre-Roman Iron Age (La Tène) artefacts and ceramics were identified from the Balve Cave youngest archaeological layers (Brandt, 1939; Bleicher, 1983, 1991).
22The studied lion bone material of the Balve and other Sauerland caves are from the collections of the Museen Burg Altena (MBA). The unlabelled and not even well-cleaned material was rediscovered within these studies. Only one hyena maxillary and about 60 teeth, less postcranial bones (mainly from the cave bear), and also several mammoth teeth, represent this Balve Cave material, which also includes about 100 Palaeolithic artefacts. Other material from C. Kremer (old labels on the bones with his name) was studied in the Goldfuss-Museum Bonn (GMB); not lion remains, but hyena and “nibbling stick” remains. The most important historical hyena and prey bone collection is from the Museum Balve (MB), which is partly on displayed and with other remains including lion material stored in the Archaeological Museum Herne. In the Emschertalmuseum Schloß Strünckede Herne (EMSH), a few bones, including hyena and lion, are deposited. A large part of the Balve Cave (coll. Kremer 1873), Martins Cave (coll. Schaaffhausen 1875; Schmitz 1889), or Wilhelms Cave (coll. Hundt and Troschel 1874) bone collections are mainly in the Geologisch-Paläontologische Museum of the Westphalian Wilhelms-University Münster (GPIM) but also in the GMB. Bones from the Keppler Cave, Hohle Stein Cave, or Johannes Cave, are housed in the Stadtmuseum Menden (SMM). Finally, a few non-lion bones (e.g. hyena) are in the collection of the Museum für Natur und Umwelt Osnabrück (MNUO). A lion jaw from the Kreuz Cave is in the Dechenhöhlenmuseum Lethmathe (DHM).
23The work was made to understand the lion bone presence at hyena den type sites from the palaeobiological point of view in the Sauerland Karst region at 13 hyena cub raising, commuting, and prey accumulation sites.
24None of the Balve Cave lion bones have exact stratigraphic context or labels with further information, but bones from older Middle Pleistocene layers are preserved different as from younger layers, and are more fossilized. The Middle Pleistocene bones generally have typical black manganese dendritic surfaces, and break due to beginning lithifications. The colour is more reddish and irregular. Late Pleistocene bones are not fossilized, have no dendritic surfaces and are more yellow to brownish in colour. Even with those helpful criteria it is too vague to present a list with their Saalien or Weichselian ages. In some cases the material might be distinguished from each other, but not in all cases, which makes again difficult a statistical analyses and identification for each bone. After the preservation degree, about 85 % and even more of the bones of Balve Cave might be expected to be from the late Pleistocene (early to middle Weichselian), and only a few seem to be from the late Middle Pleistocene (late Saalian). This generally is coincident to the amount of all bone material (mostly from early to late Weichselian), which is barely represented in the oldest layers. The Saalian horizons were historically not much excavated, because they often did not reach deep or even the cave base. The stratigraphic information about the finds is therefore limited. As demonstrated in the stratigraphy, there are three late Pleistocene horizons which must have produced the here-described lion material, whereas in the historical description, Panthera bones are only mentioned for the early Weichselian layers.
25With the historical lion bone material from the Sauerland Karst at least sex identification was tried for complete bones, whereas an individual age statistics is the most valuable information, such as taphonomic studies including bone damages and bite marks caused by carnivores. Also the bone frequency to estimate the body part presence is presented, but the material from all caves again is too few for a good data base.
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Family Felidae Gray 1821
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Genus Panthera Oken 1816
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Panthera leo spelaea (Goldfuss 1810)
26(fig. 4, 5, 6, 7, 8, 9 & 10)
Fig. 3: Panthera leo spelaea (Goldfuss 1810) skeleton part remains from the balve cave and other caves of the sauerland Karst Mountains (NW Germany), all being from cave bear or hyena dens.
Fig. 4: Panthera leo spelaea (Goldfuss 1810) cranial remains from the Balve Cave (NW Germany).
1. Brain case from a young adult lioness (MB No. Balve-1): a. lateral right, b. occipital, c. dorsal, d. ventral. 2. Brain case from an old adult to senile lion (MB No. Balve-2): a. lateral right view, b. occipital view, c. dorsal view, d. ventral view. 3. Left upper jaw canine from an adult individual (GPIM No. A5F1297): a. caudal view, b. labial view. 4. Right lower jaw canine from an adult animal (MB No. Balve-4): labial view. 5. Left mandible from a lioness (MB No. 861): lateral left view.6. Mandible fragment from a lioness (MB No. Balve-3): lateral right view. 7. Left mandible fragment (EMSH No. Bal-1): lateral view. 8. Right upper jaw P3 from an adult animal (GPIM No. A5F1265): labial view. 9. Left lower jaw M1 from a senile animal (GPIM No. A5F1296): labial view.
Tab. 1: Panthera leo spelaea (Goldfuss 1810) bone remains from the Balve Cave (NW Germany).
Fig. 5: Panthera leo spelaea (Goldfuss 1810) vertebra, fore and hind limb bone remains from the Balve Cave (NW Germany). 1. Cervical vertebra
No. 5 (GPIM No. A51022): a. dorsal view, b. cranial view. 2. Right mc IV and V from a lioness (MB No. Balve-7/8): dorsal view. 3. Right mc V of a young adult animal (MB No. Balve-10): dorsal view. 4. Left mc V from an adult lion (MB No. Balve-11): dorsal view. 5. Left mc II from an adult lion (MB No. Balve-12): dorsal view. 6. Right scapholunatum (GPIM No. A51023): dorsal view. 7. Right capitatum (GPIM No. A51024): cranial view. 8. Left tibia from an adult lion (MB No. Balve-14): cranial view. 9. Left astragal from an adult lion (MB No. Balve-15): dorsal view. 10. Left astragal from an adult lioness (MB No. Balve-18): dorsal view. 11. Left cuboid (GPIM No. A51025): cranial view. 12. Left mt V of an adult lion (MB No. Balve-23): dorsal view. 13. Left mt V from an adult lion (MB No. Balve-24): dorsal view. 14. Right mt IV and V from an adult lioness (MB No. Balve-21/22): dorsal view. 15. Right mt IV from an adult lioness (MB No. Balve-30): dorsal view. 16. Right mt IV from an adult lioness (MB No. Balve-43): dorsal view. 17. Left mt IV of an adult lion (MB No. Balve-28): dorsal view. 18. Right mt III from an adult lion (MB No. Balve-29): dorsal view. 19. Right mt III from an adult lion (MB No. Balve-31): dorsal view. 20. Half phalanx I digit I (MB No. Balve-36): dorsal view. 21. Sesamoid (GPIM No. A51026): lateral view. 22. Sesamoid (GPIM No. A51027): lateral view. 23. Phalanx I right digit IV from the manus (MB No. Balve-37): dorsal view. 24. Phalanx I left digit IV from the pes (MB No. Balve-38): dorsal view. 25. Phalanx I left digit III from the pes (MB No. Balve-39): dorsal view. 26. Phalanx I left digit IV from the manus (MB No. Balve-40): dorsal view. 27. Phalanx I right digit III from the manus (GPIM No. A5F1182): dorsal view. 28. Phalanx I left digit V from the manus (GPIM No. A5F1184): dorsal view. 29. Phalanx I right digit II from the manus (MB No. Balve-41): dorsal view. 30. Phalanx I left digit IV from the manus (GPIM No. A5F1183): dorsal view. 31. Phalanx I left digit V from the pes (MB No. Balve-42): dorsal view. 32. Phalanx
Fig. 6: Panthera leo spelaea (Goldfuss 1810) remains from different Sauerland Karst Mountain caves (NW Germany).
1. Right mandible from the Kreuz Cave (DHM No. Kreuz-1): lateral view. 2. Lower jaw canine from the Grürmanns Cave (GPIM No. A5F1275): lateral view. 3. P tooth fragmentfrom the Martins Cave (GPIM No. A5F1276): lateral view. 4. Left cracked humerus of a lioness from the Teufelskammer Cave (GMB No. M5832): cranial view. 5. Left humerus shaft from the Hohle Stein Cave (SMM No. 0/5523): cranial view. 6. Right cracked ulna from the Martins Cave (GPIM No. A5F1231a): a-b. lateral view, c. cranial view. 7. Right mc IV from the Johannes Cave (SMM No. 90/6528): dorsal view. 8. Left mc II from the Martins Cave (GMB No. M2466): dorsal view. 9. Lumbar vertebra no. 6 from the Johannes Cave (SMM No. 90/6390): lateral view. 10. Upper caudal vertebra from the Grürmanns Cave (GPIM No. A5F1274): lateral view. 11. Middle caudal vertebra from the Martins Cave (GPIM No. A5F1196): lateral view. 12. Right calcaneus (GPIM No. A5F1231): cranial view. 13. Right astragalus from the Martins cave (GMB No. M2460): dorsal view. 14. Right mt V from the Martins cave (GPIM No. A5F1231): cranial view. 15. Left mt IV from the Martins cave (GBM No. M2462): dorsal view. 16. Right mt IV from the Martins cave (GBM No. M2470a): dorsal view. 17. Right mt III from the Martins Cave (GMB No. M2470): dorsal view. 18. Right mt II from the Martins cave (GMB No. M2463): dorsal view. 19. Rigth mt II from the Martins cave (GMB No. M2464): dorsal view. 20. Phalanx I from the Martins Cave (GPIM No. A5F1185): dorsal view. 21. Phalanx I from the Martins Cave (GPIM No. A5F1197): dorsal view. 22. Phalanx I from the Martins Cave (GPIM No. A5F1198): dorsal view.
Fig. 7: Panthera leo spelaea (Goldfuss 1810) cranial and fore limb remains from Keppler Cave of the Sauerland Karst Mountains (NW Germany).
1. Right mandible of an adult lioness (SKMM No. 90/7047): lateral view. 2. Left mandible of an adult female (SKMM No. 90/7046): lateral view. 3. Left mandible of an adult male (SKMM No. 90/6495): lateral view. 4. Right M1 of an adult lioness (SKMM No. 90/7048a): labial view. 5. Left M1 of an adult lioness (SKMM No. 90/7048b): labial view. 6. Right humerus of an adult to senile male (SKMM No. 90/6521): cranial view. 7. Right humerus of an adult to senile lioness (SKMM No. 90/6520): a. cranial view, b. lateral view, right with bite scratches. 8. Right humerus of an adult to senile male (SKMM No. 90/6522): cranial view. 9. Left Mc IV of an adult to senile lioness (SKMM no. 90/6531): cranial view. 10. Left Mc IV of an adult to senile lioness (SKMM No. 90/6528): cranial view.
Fig. 8: Panthera leo spelaea (Goldfuss 1810) vertebral column remains from Keppler Cave of the Sauerland Karst Mountains (NW Germany).
1 Axes of an adult animal (SKMM No. 90/6273): a. cranial view, b. lateral left view. 2. C3 of an adult animal (SKMM No. 90/6799): a. cranial view, b. lateral left view. 3. C5 of an adult animal (SKMM No. 91/8751): a. cranial view, b. lateral left view. 4. C6 of an adult animal (SKMM No. 90/6754): a. cranial view, b. lateral left view. 5. T1 of an adult animal (SKMM No. 91/8752): a. cranial view, b. lateral left view. 6. T2 of an adult animal (SKMM No. 91/8753): a. cranial view, b. lateral left view. 7. T4 of an adult animal (SKMM No. 90/6795) a. cranial view, b. lateral left view. 8. T7 of an adult animal (SKMM No. 90/6806) a. cranial view, b. lateral left view. 9. T8 of an adult female (SKMM No. 91/8754) a. cranial view, b. lateral left view. 10.T9 of an adult animal (SKMM No. 91/8755) a. cranial view, b. lateral left view. 11. T10 of an adult animal (SKMM No. 91/8756) a. cranial view, b.lateral left view. 12. T14 of an adult animal (SKMM No. 91/8757): a. cranial view, b. lateral left view. 13. L3 of an adult animal (SKMM No. 90/6750): a. cranial view, b. lateral left view. 14. L5 of an adult animal (SKMM No. 91/8758): a. cranial view, b. lateral left view. 15. L5 of an adult animal (SKMM No. 91/8759): a. cranial view, b. lateral left view. 16. L6 of an adult animal (SKMM No. 91/8760): a. cranial view, b. lateral left view.
Tab. 2: Bone list of Panthera leo spelaea (Goldfuss 1810) remains from the Keppler Cave in the Sauerland Karst Mountains (NW Germany) from the Heinrich 1919-1920 collection
Fig. 9: Panthera leo spelaea (Goldfuss 1810) hind limb remains from Keppler Cave of the Sauerland Karst Mountains (NW Germany).
1. Left pelvic fragment of an adult to senile animal (SKMM No. 90/6962): lateral view. 2. Right femur of an adult to old lion with scavenged marks on the distal joint (SKMM No. 90/6357): a-c. cranial view. 3. Right femur of an adult to senile lioness (SKMM No. 90/6524): cranial view. 4. Left tibia of an adult to senile male (SKMM No. 90/6525): cranial view. 5. Right Mt II of an adult to senile lioness (SKMM No. 90/6526): cranial view. 6. Right Mt III of an adult to senile lioness with a chewed distal joint (SKMM No. 90/6527): a-b. cranial view. 7. Left Mt III of an adult to senile male (SKMM No. 90/6530): cranial view.
Tab. 3: Panthera leo spelaea (Goldfuss 1810) bone remains from the Martins Cave (NW Germany).
Fig. 10: Cub skeleton Panthera leo spelaea (Goldfuss 1810) remain from the Wilhelms Cave hyena den (NW Germany).
1. Right upper jaw canine (GPIM No. A5F1179): labial view. 2. Right scapula (GPIM No. A5102-1): lateral view. 3. Left scapula fragment (GPIM No. A5102-2): lateral view. 4. Left humerus (GMB No. M6006): cranial view. 5. Right humerus (GPIM No. A5F1180): cranial view. 6. Axes (GPIM No. A5F1298): a. lateral left view, b. dorsal view. 7. Last lumbar vertebra L7 (GPIM No. A5F1299): lateral right view, b. cranial view. 8. Sacrum (GPIM No. A5102-3): a. cranial view, b. dorsal view. 9. Middle left costa (GPIM No. A5F1225): caudal view. 10. Middle right costa (GPIM No. A5102-5): caudal view. 11. Middle right costa (GPIM No. A5F1088): caudal view. 12. Right femur shaft (GPIM No. A5102-4): cranial view. 13. Right calcaneus (GPIM No. A5F1262): cranial view. 14. Left calcaneus (GPIM No. A5F1263): cranial view. 15. Right astragal (GMB No. M792): dorsal view.
27Two brain cases of incomplete skulls, a few incomplete mandibles, some isolated teeth, one complete tibia, astragals and calcanei, but mainly metapodials and phalangae; in total 56 bones (tab. 1). Most of the material is not strongly fossilized and most probably mainly from the early to middle Upper Pleistocene layers. Only main measurements are given for possible comparative sex identifications.
28The first skull is, in its preservation, typical of late Pleistocene bone material. This one is small in proportions and consists only of the brain case (fig. 4.1). The cranial bones are not well-fused. The sagittal crest is already developed indicating an age of an older juvenile to a young adult animal. The measurements of the condyle width (61 mm) and frontals width (81 mm) would match young female skulls too. A total length of the brain case of 192 mm can be given with an estimated short total length of about 300 mm. The posterior brain case damage is similar to the damage on the second skull; this is old damage that seems to be the result of carnivore activities, such as hyenas scavenging damage. The dentition is absent.
29The second skull (fig. 4.2) is much larger in its brain case proportions compared to the first skull. Again, the saggital area is missing parts. The anterior brain case has modern fractures; originally the skull possibly had its maxillaries. The occipital width (69 mm) is large and fits to grown up and male skulls as discussed later. The total preserved length is 170 mm. As a result of its completely fused brain case sutures, the individual was an older adult.
30A single maxillary canine from an adult individual is present (fig. 4.3). The root is completely closed and the sharp edges are not rubbed, supporting an adult individual age. Only one lower jaw canine, which is slightly incomplete in its enamel area (fig. 4.4), is from an adult animal with small proportions.
31The lower jaws are all incomplete. The most complete left one (fig. 4.5) lacks the ramus due to modern damage, but has the P3-4 and M1 teeth. The width of the molar cannot be measured, as it was proximally broken during the lifetime of this old adult animal, and slightly polished. The P4 length is 29 mm. A second right mandible (fig. 4.6) is represented by its anterior part with half of the P3 and the P4. The small P4 length is 29 mm. The third mandible is a fragment including the P4 (fig. 4.7). Only two teeth, a complete right upper P3 (length 30 mm) from an adult animal (fig. 4.8), and a left lower M1 without parts of the root (fig. 4.9), from a senile one, are preserved in the material.
32The fore limbs are missing all long bones. This is most likely a problem of historical collecting, material selection and loss. At least seven metacarpi and six first phalangae are preserved. The gender of the metacarpi can be possibly determined by its proportions; however it is impossible to make this determination for the phalangae. The metarcarpi (fig. 5.2, 5.3, 5.4 & 5.5), all from adult individuals, are listed in table 1 with main data and most probable sex identifications.
33Important are the two metacarpi that fit (fig. 5.2) because of their similar preservations and small proportions. This indicates the presence of originally articulated pedal skeletons at the cave site. Strong different proportions are not only visible in the two mc V (fig. 5.3 & 5.4). The phalangae (fig. 5.23-33), are from different digit positions of the manus (see also tab. 1).
34The hind limbs are represented at least by one long bone, a left tibia (fig. 5.8), from an adult male lion. Thirteen metatarsi, all from adult animals are the dominant bone types - all data are in table 1. To mention are the different proportions (cf. fig. 5) and two well-matching right metatarsi IV/V (fig. 5.14). The latter ones prove articulated pedal skeletons at the cave site. The phalangae I are well-represented in the material; their digit position can mostly be given. Manus and pes phalanges are from different positions as mentioned (see tab. 1, fig. 5.23 to 33).
35The Upper Pleistocene Bilstein Cave lion material from this cave bear den (show cave) with short hyena occupation in the entrance area was monographically figured and described (Diedrich, 2009d). 39 bones are present from all body parts of different male and female individuals; all adult are old in age. Only one bone is from a young adult animal.
36A single modern broken canine of an adult animal is the first remain from this hyena den cave (fig. 7.2); the second is an upper caudal vertebra (fig. 7.10).
37From this destroyed small hyena prey depot cave at the beginning of the Hönnetal, a single small-sized and modern damaged shaft, probably from an early adult animal (fig. 6.5), is the only lion remain.
38Also this small cave destroyed around 1944 was a hyena den. Lion remains are a metapod (fig. 6.7), and a lumbar vertebra (fig. 6.9), both remains from grown up individuals.
39The Keppler Cave in the Hönnetal was opposite to the Balve Cave and is completely destroyed today. In total, 33 bones are preserved in the large cave bear dominated bone collection (fig. 7, 8 & 9), which show, in some cases, modern fractures as a result of the historical excavations. Three mandibles and two isolated lower jaw teeth (M1) are the cranial remains. The first right lower jaw (fig. 7.1) has modern fractures all over. The P3-4 and M1 teeth are slightly rubbed, indicating an adult animal age. The width of the M1 (29 mm) is quite small. The second left mandible (fig. 7.2) has no ramus and only the P3-4 and M1 teeth. The latter posterior tip was broken during the animal life, indicating an older adult age by its tooth wear. The small proportions, such as the mandible height between the P4/M1 (47 mm) are slender; also the M1 is only 24 mm in length. The third left mandible (fig. 7.3) is strongly encrusted by a speleothem layer. Only the P4/M1 are preserved; the molar has old fractured tips indicating an older adult individual. The two isolated teeth from adults are both M1 (right and left) and each is 27 mm in length.
40The fore limb bones are three right humeri (fig. 7.6, 7.7 & 7.8); two of them are very long (360 and 362 mm), only one is much shorter (342 mm length). All are adult to older individual remains. The lioness humerus has 2-3 mm thin bite scratch marks on the shaft (fig. 7.7b). Two left metacarpi IV (fig. 7.9 & 7.10) are again from adult to old animals; they are short in their lengths (117/122 mm).
41A large amount of 16 vertebrae consists of four cervical, eight thoracic and four lumbar vertebra, all generally missing parts of the processi because of modern fractures. The second cervical vertebra, the axes (fig. 8.1), the third (fig. 8.2), the fifth (fig. 8.3) and the sixth (fig. 8.4) are all quite small in size. They might even belong to one skeleton. Also, the thoracic vertebrae seem to not be represented twice. The T1 (fig. 8.5) is quite small and, whereas the T2 (fig. 8.6) is large. There are four more middle to posterior thoracic small-proportioned vertebrae; a T4 (fig. 8.7), T7 (fig. 8.8), T8 (fig. 8.9), T9 (fig. 8.10), T10 (fig. 8.11), and finally a complete T14 (fig. 8.12). Four lumbar vertebra consist of one L3 (fig. 8.13), whereas one L5 is also smaller (fig. 8.14). The other similar L5 is larger (fig. 8.15). Finally a sixth lumbar vertebra (fig. 8.16) is again smaller sized.
42One pelvic fragment from the left side (fig. 9.1) is from an adult animal, whereas the acetabulum diameter with is 49 mm. Hind limb bones are present, all from adult animals, with two femora, one tibia and three metatarsi. The first right and distally larger male femur is proximally modern broken and chewed distally by carnivores (fig. 9.2a-c). One complete right femur is most likely from a large animal (fig. 9.3) with its 405 mm length and distal width of 60 mm. A complete left tibia (fig. 9.4) measures 355 mm in length and falls into the female size-range. The pedal skeleton is represented by a right mt II (fig. 9.5); a right mt III from with a chewed distal joint (fig. 9.6), and finally a left mt III from an adult to older animal (fig. 9.7).
43A right mandible of a grown up individual was found in unclear context and as single remain in the cave (fig. 7.1).
44The 17 bones were excavated in the hyena den cave near Lethmathe (Schaaffhausen, 1875) which are listed in table 3, and are figured here in most cases. A half P tooth (fig. 7.3) is the only cranial remain. The fore limb is preserved by a cracked and chewed ulna of a grown up animal (fig. 7.6), and several metacarpi (fig. 7.7 & 7.8). The axial skeelton has one incomplete middel caudal (fig. 7.11) and one lumbar vertebra preserved. The astragalus has bite damages (fig. 7.12), one astragalus is complete (fig. 7.13). The hind limb is represented additionally by five metatarsi (fig. 7.14, 7.15, 7.16, 7.17, 7.18 & 7.19). The phalany I bones (fig. 7.20, 7.21 & 7.22) are unclear in their pedal positions, but one has bite damages.
45The only nearly complete skull of a lioness was described and all figured monographically with a taphonomic discussion of the remains in a cave bear and hyena den (Diedrich, 2008b).
46From this hyena den, only a non-figured metapodial and one phalanx II are in the few today preserved historical collection.
47This cave was a hyena den only during the Upper Pleistocene, for which an individual skeleton remain of a young lion cub found in historic times (Hundt & Troschel, 1874) is remarkable (fig. 10). 15 bones are preserved, which show in some cases modern fractures. One upper jaw right canine is open from the root (fig. 10.1). From the fore limb both incomplete scapulae (fig. 10.2 & 10.3) are preserved, of which the left one consists only of the proximal part. The height of the right scapula is 222 mm. The right humerus measures 271 mm in length without its proximal joint. Both humeri are lacking its proximal joints. The right one has a fused distal joint, giving an individual age of a cub. The left humerus distal joint was broken modern. The vertebral column is represented by the axes (fig. 10.6) and the last lumbar vertebra No. 7 (fig. 10.7); both are missing the centrum discs. From the pelvic bone, the sacrum (width of 78 mm with the first two elements: fig. 10.8) is incomplete. Also here the disc is missing. The rib cage is represented by three incomplete, middle thoracic ribs, two of them are from the right side (fig. 10.10 & 10.11), one costa is from the left thorax (fig. 10.9). The joint head discs are absent and were non-fused. The here-figured lion ribs were formerly mounted incorrectly in a hyena skeleton montage consisting mainly of hyena bones from this cave (Humpohl et al. (1997); there named “Biggethal Cave”). There are four hind limb bones: one right femur shaft 292 mm in length (fig. 10.12), both calcanei (length 106 mm each), without distal parts (fig. 10.13 & 10.14) and the right astragal (length 58 mm) are preserved (fig. 10.15).
48An incomplete small proportioned humerus from an early adult animal, which was cracked in the upper third of the shaft and was chewed laterally at the distal joint by a large carnivore (fig. 7.4).
49The lion material was compared with other descriptions of European lions of the Middle to mainly Upper Pleistocene periods (cf. Heller, 1953; Dietrich, 1968; Schütt, 1969; Altuna, 1981; Turner, 1984; Argant, 1988; Gross, 1992; Guzvica, 1998; Baryshnikov and Boeskorov, 2001; Burger et al., 2004; Diedrich, 2004, 2007b, 2008a, 2009a, 2009c, 2009d, 2010b, 2010c, 2011c; Diedrich & Rathgeber, accepted). Some metapodials from the Balve Cave are extremely large and seem to fit more to maximum-sized Saalian steppe lion remains compared to the descriptions of the few record of European material from the late Middle Pleistocene (Argant, 1988).
50Anyway, here some of the Upper Pleistocene lion bones are compared in their proportions to other Middle European studied caves in Germany, and Czech Republic. Included are also a few data of Gross (1992), for which metric scheme was used. The few represented long-bone material of the Balve Cave was recently compared in another study of the Bilstein Cave, and Keppler Cave lions, two cave bear den sites in the Sauerland Karst without any human impact. Those caves are not far from the Balve Cave (see fig. 1). Also the Perick Cave material was used, which instead is from a cave bear and hyena den site. The lion long-bone data are not repeated here, because there is only one tibia in the Balve Cave material, which falls into the range of larger males (Diedrich, 2009d). Instead, other postcranial bones are compared statistically with Middle European caves, gypsum karst and open air hyena den sites by the calcaneus and astragalus (fig. 6A & 6B). The astragals of the Balve Cave might indicate a sexual dimorphism, whereas at least three of them seem to be smaller female ones, if all would be from cold periods and not from Eemian smaller sized proportioned lions (Eemian lioness skeleton: Diedrich, 2010c) or if those are not from larger sized late Middle Pleistocene lionesses (Argant, 1988). One is in the uppermost range of males compared to cold period Weichselian/Würmian large lions (fig. 6B).
51The lion material from the Balve Cave is not represented well enough to perform more detailed analyses. However, the material is still one of the four largest lion bone collections of the Sauerland Karst in Westphalia; compared to the Perick Caves, Keppler Cave, Bilstein Caves, Wilhelms Cave and others (Diedrich, 2009a, 2009b, 2009d), and the only larger lion bone collection from a human influenced cave site of this region (sites: fig. 1). The early to middle Weichselian lion material of the Perick Caves hyena and cave bear den contain the only known, nearly complete, lioness skull (Diedrich, 2009a) of the Sauerland Karst. The Wilhelms Cave has produced skeleton remains from a lion cub, which hyenas must have been imported as kill into their den cave. As described for the Bilstein Caves (Diedrich, 2009d); the Keppler Cave herein has also delivered only adult lion material. All lion remains occur at all cave sites, that are hyena den caves and cave bear dens (cf. Diedrich, 2009b). Additionally to the material from other German and Czech localities, other material was integrated from the hyena den gypsum karst sites Thiede (Lower Saxony), Quedlinburg Sevecken-Berge and Westeregeln (Saxony-Anhalt), or material from the Zoolithen Cave (Bavaria), such as the skeleton find from the Czech Srbsko Cave (Bohemian Karst), and material from the Sloup and Výpustek Caves (Moravian Karst). Finally, from the Westphalian open air site Bottrop (fig. 1) proportions of bones were compared (fig. 11).
52Eighteen female bones (58 %) and 13 male bones (42 %) can be identified in the Balve Cave material due to their noticeable size differences. All other bones are too incomplete, or represent intermediate sizes, which are difficult to use when attempting to identify the sex. In the Bilstein Cave, the lioness material (NISP = 38 bones; Diedrich, 2009d) dominates with 67 % (33 % males). Keppler Cave lion material includes at least two lion and two lioness remains, all being from adult to senile individuals. The percentage of lionesses seems to be, with 22 bones (67 %) higher then lion remains (11 bones = 33 %). Juvenile or adolescent material is absent. In the Perick Caves, it is opposite, here 66 % are males and the rest are from females (NISP = 59 bones; Diedrich, 2009a). More caves, and especially hyena dens, must be studied in their lion bone content to form a theory about possible male or female lion dominance in cave sites.
53
Fig. 11. Panthera leo spelaea (Goldfuss 1810) sexual dimorphism comparisons of the hind limb bones.
A. Calcaneus and B. Astragalus of German and Czech caves, gypsum karst and open air sites, nearly all identified as hyena dens (long bone comparisons in Diedrich (2009d); data in cm).
Fig. 12: comparisons of Panthera leo spelaea (Goldfuss 1810) percentages (1-4 %) and carcass destruction stages in the most important non-human influenced sauerland Karst hyena and cave bear den caves.
The faunal statistics presents typical cave bear den and hyena den bone assemblages. The faunal percentages depend on the hyena impact and hyena prey importation.
54General human versus carnivore bone accumulation studies were tried to distinguish bone assemblages using actuopalaeontological comparisons (e.g. Cooper, 1993; Fosse, 1999; Pickering, 2002; Stiner, 2004; Lansing et al., 2007). New excavations at Balve Cave in future must focus on such taphonomic problems with the background knowledge of the multiple cave use by hyenas, cave bears and humans.
55The Balve Cave was recently identified as a periodically used hyena den for at least the early to middle Weichselian period (Diedrich, 2011a). The criteria for the late Pleistocene hyena den and communal site identifications are followed sensu Fosse (1999), Diedrich and Žák (2006), and Diedrich (2011d). After Kuhn et al. (2008) in modern hyena “accumulated bone assemblages” a high amount of bones with crenulated edges and punctures are present. This is also observed on the selected “incomplete bone material” of the Balve Cave, which can be referred in many cases well to Ice Age spotted hyena activities (Diedrich, 2011a). Another proof for a hyena den and commuting site at the Balve Cave is a relatively large amount of hyena Crocuta crocuta spelaea bone material including cannibalistic cracked hyena bones or jaws and cub remains or imported, strongly chewed megafaunal prey remains, especially from Coelodonta antiquitatis (Diedrich, 2011a). The woolly rhinoceros bones are in most cases strongly damaged in repeating ways as figured similar for many other hyena den and prey accumulation sites: Bad Wildungen Biedensteg open air hyena den and prey depot (Diedrich, 2006), Perick Caves hyena den and prey depot (Diedrich, 2005, 2009a), and several hyena den and prey depot caves in the Bohemian Karst (Diedrich & Žák, 2006). It is less the numeric analyses that support the hyena origin to distinguish from human sites; it is more the repeating similar incomplete bone damage including “nibbling sticks” (cf. Diedrich, 2006, 2011a; Diedrich & Žák, 2006).
56As established, juvenile hyena remains of the Balve Cave (Diedrich, 2011a) are best indicators for “hyena bone accumulation proves” (cf. Kuhn et al. 2008), they are accepted well to be the indicators for hyena den sites (cf. Pickering, 2002). Those are recently known for the Pleistocene to indicate even the den type of a “cub raising den” (cf. modern: East et al., 1989; Pleistocene: Diedrich, 2006, 2011a; Diedrich & Žák, 2006). Such modern cub raising and protecting den sites are well-known for African spotted hyenas (e.g. Kruuk, 1972; Cooper, 1993; Frank, 1994; Estes, 1999). In contrast to those cub dens, their remains are mostly absent at “commuting or prey accumulation sites” (definitions for Pleistocene dens sensu Diedrich & Žák, 2006). Hyena remains are sometimes absent even at hyenid “bone accumulation sites” (cf. Kuhn et al., 2008).
57Pickering (2002) and Kuhn et al. (2008) suggested in modern African spotted hyena bone assemblage analyses that horns and antlers are not indicative for hyena dens, which has to be discussed shortly for the Balve Cave antler remains. Hyenas collected dropped antlers. Humans also did it, but the antler preservations is different. Humans collected those for tool productions, therefore for more then 50 reindeer antler fragments of selective mainly larger male ones were found in the Balve Cave with no bite marks. It is less the antler presence in the hyena dens that indicates a hyena origin, than the repeated chewed basal remains hyena left also in the Balve Cave with some specimens of Rangifer tarandus and Megaloceros giganteus antler bases (Diedrich, 2011a). Wolves do not import such dropped antlers to caves, or do not chew strongly on them, which was also excluded for the Italian hyena den caves (Stiner, 2004).
58About 250 lion bones of the Sauerland Karst Caves of north-western Germany are dominated by adult to senile lion remains (99 %). This supports the theory of the non-use of caves by lions as a “den” to raise their cubs in caves or cavities. They seem never to have used caves, and their occurrence in those caves is partly a result of the antagonism between Ice Age spotted hyenas and steppe lions (Diedrich, 2007a; in press), but also a result of cave bear killing and scavenging by lions (fig. 7), which seem to be present also in Keppler Cave cave bear den. Therefore, the lions were renamed from “cave lions” to “steppe lions” (Diedrich, 2008a).
59Most lion remains in Balve Cave are from the hyena den times of the late Pleistocene. The import of lion carcasses into hyena den sites is not only suggested for the Sauerland Karst caves (Diedrich, 2009a), it is discussed for some Bohemian Karst hyena den and prey depot caves (Diedrich & Žák, 2006). The presence of many lion metapodials is common at several studied hyena den sites and can be explained partly by the selective importation of legs. Other lion bones were damaged by hyenas in the main eating zones around the scapula/humerus and pelvic/femur areas, as recently described for the lion material from the Perick Caves (Diedrich, 2009a; fig. 3). Hyenas mostly did not scavenge on pedal skeletons from any kind of megafaunal animal leaving hereby within ungulate prey, the lower legs remaining untouched in their dens (cf. Stiner, 2004; Diedrich & Žák, 2006; Diedrich, 2010a), but there are exceptions. The distal pedal parts of carcasses were mostly left from the imported prey legs, because of the small amount of meat and missing bone marrow (cf. Stiner, 2004; Diedrich, 2010a). Typically, hyenas leave at lion scavenge sites the cranial bones, lumbar vertebrae, distal or complete legs, such as those described for Perick Caves (Diedrich, 2009a). In Perick Caves, the lion bones were in several cases cracked (lower jaws), nibbled and chewed; similar strongly damaged are bones damaged from Martins Cave hyena den. Nearly no scavenging marks were found on the lion bones from Balve Cave, which is in contrast to the lion bones from Martins Cave, Keppler Cave, Bilstein Cave, and Perick Caves (cf. Diedrich, 2009a, 2009d, 2011d), where several bones have bite marks caused by a large predator, most probably hyenas. At least there are some elongated parallel 3-4 mm wide bite scratch marks on Balve Cave tibia, caused by a large carnivore. Possibly both occipital areas of Balve Cave skulls are missing parts because of carnivore activities; this is difficult to prove, but at least it is another proof of hyena activity. A similar preservation was found at a lion brain case of Perick Caves (Diedrich, 2009a).
60The large Sauerland caves were occupied by two other large animals, hyenas and cave bears. Late Pleistocene lions were open steppe and forest predators, similar to their modern relatives in Africa (Diedrich, 2007a, 2009a) and seem to have penetrated for two reasons the caves: (1) stealing hyenas their prey at the commuting or prey storage sites; or to kill their cubs in the raising cave sites as result of antagonism, and (2) for cave bear hunting (Diedrich, 2009a). In both cases antagonistic conflicts must be expected which caused the death of lions directly in caves. The Balve Cave, and the Keppler Cave material especially supports both theories that lions were killed by hyenas and possibly by cave bears in the caves, whereas Martins Cave material seem to have been imported by hyenas as carcass remains mainly (cf. Diedrich, 2009a).
61In modern studies, information about lion or other large carnivore bone remains at hyena bone accumulations is poor. Only Lam (1992) and Pokines and Peterhans (2007) mentioned for few modern spotted hyena den sites a comparative proof for the importation of lion carcass remains. For the late Pleistocene studied bone-rich cave and open air sites, this possibility why steppe lions were found in European caves or at hyena dens coincidently was discussed (Diedrich, 2007a, 2007b, 2008a, 2009a, 2009b, 2009d), but new articulated skeleton lion finds in a cave in Romania (Diedrich et al., 2009) proove again their presence even very deep in cave bear den caves, as most probable cave bear hunt conflict result.
62It is known that modern African spotted hyenas do kill usually the weaker lionesses, but never the male lions (Sutcliffe, 1970; Estes, 1999; Joubert & Joubert, 2003). Those kills are for antagonistic reasons (Ford, 2005). Hyenas also possibly imported carcasses of dead lions that died a natural death or from intra-species fighting (Schaller, 1972; Palomares & Caro, 1999).
63The comparison of lion remains at non-human influenced sites (Perick Caves, Keppler Cave, Bilstein Cave, Wilhelms Cave, and others in the Sauerland Karst (cf. fig. 1, 12) excludes evidence, even here in the Balve Cave, of the import of lions by humans. There are no cut marks at the Balve Cave on the lion bones, especially the most common metapods. A large amount of manus and pes skeleton elements is not proof of a lion “fur import by humans” following possible lion kills by humans. This taphonomic situation is important, because there still remains the question if humans hunted, killed and imported lions to their settlement and camp sites in the Middle Palaeolithic. Only the presence of lion bones does not prove this. This is demonstrated here for the Balve Cave; where hyena activity is responsible for the large amount of accumulated bones, similar to the non-human influenced hyena den Perick Caves (Diedrich, 2009a) or Srbsko Chlum Komin Cave (Diedrich & Žák, 2006). It is not yet proven whether lions or hyenas were hunted and killed by humans; both of which were the most dangerous predators and antagonists for humans and to each others and to cave bears at that time in Central Europe, but conflicts also have to be expected.
64
Fig. 13: Panthera leo spelaea (Goldfuss 1810) male with mane and typical tassel on the tail tip, and lioness in the background, in boreal forest environments of the Sauerland Karst Mountains
Illustration by G. “Rinaldino” Teichmann, 2010
65The lion Panthera leo spelaea (Goldfuss 1810) bone material presence in all northwest German Sauerland Caves originates of two main carnivore ecological situations, and can be excluded in the overlapping Balve Cave Palaeolithic site to be “imported prey of humans for meat or fur purposes”. The Balve Cave was several times used as periodic hyena Crocuta crocuta spelaea (Goldfuss 1823) den. Therefore some remains of lions, which have even bite damages, might reflect imported and consumed carcass remains by hyenas, such as demonstrated for the Perick Caves hyena den lion remains. Additionally, lions might have been killed at the hyena den Balve Cave site due to antagonistic conflicts about prey or cub protection or territory. Also lion-cub-killing and import of its carcass seem to be present at the Wilhelms Cave hyena cub raising den site. Finally, supporting the “lion-cave bear killing model” for European caves, as a result of the mammoth steppe fauna scarcity/absence in mountainous regions, lion remains are typically found in few percentages (1-3 %) in nearly all cave bear dens of Europe, such as the German Zoolithen Cave, Bilstein Cave, the Keppler Cave and the periodically used cave bear den Balve Cave.
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