Crocuta KAUP 1828

GENUS <i>CROCUTA</i> KAUP, 1828 <p> <i>CROCUTA CROCUTA SPELAEA</i> (GOLDFUSS, 1823)</p> <p>(FIGS 2, 3)</p> <p>1810 ‘Hölen Hyäne’. – Goldfuss: 280–281, table V, figs 2a, b.</p> <p> 1823 <i>Hle</i>. LVI, figs 1–3.</p> <p> <i>Holotype:</i> <i>Hyaena spelaea</i> Goldfuss, 1823. Incomplete cranium (Goldfuss-Museum Bonn No. M2609).</p> <p> <i>Locus typicus:</i> Zoolithen Cave, Geilenreuth (cave No. D 106) Franconian Alb, Wiensent Valley, Bavaria, southern Germany, Central Europe.</p> <p> <i>Stratigraphy:</i> ‘Upper layers above the main cave bear bone gravels’ after Goldfuss (1810, 1823). Weichselian/Würmian, Upper Pleistocene (probably OIS 3). OIS 3–6 after Rosendahl & Kempe (2004). A radiocarbon date on cave bear teeth is given as 28.905 ± 255 BP (Poll, 1972).</p> <p> <i>Collection:</i> Goldfuss-Museum of the Rheinische Friedrich Wilhelms-Universität Bonn, Germany.</p> <p> <i>Short description and discussion:</i> The holotype was described as an incomplete cranium (Fig. 2A), which is missing the left maxilla and premaxilla (cf. Goldfuss, 1810). The right maxilla has the alveoli of the P 3–4. The P 2 is represented by its roots and indicates a mature to very old individual. The gender of the hyena is not yet clear as a result of lack of premaxillaries and total length measurements. The Late Pleistocene hyenas often show clear sexual dimorphism, clan-leading females being slightly larger with skull length of 30–32 cm (e.g. Diedrich, 2005, 2007b); the general sexual dimorphism in <i>C. c. spelaea</i> is present in the modern African spotted hyena <i>C. c. crocuta</i> (cf. Rohland, 2003). Recently, parts of the saggital crest have been damaged (compare Figs 2 and 3A); this damage was not present in 1810, when the skull was figured in a lithography. One additionally figured lower jaw in 1810 could not be found recently and it be shown to belong to the skull. Many mandibles were found in this cave, and it is thus not possible to prove that both parts are from one individual. In contrast, the skull must be from a very old animal. The P 2 is represented by rounded roots; both M 1 s are missing and their alveoli were already completely closed. The historically figured mandible was from an adult animal and has much less worn teeth.</p> <p>FAMILY FELIDAE FISCHER, 1817</p> <p> GENUS <i>PANTHERA</i> OKEN, 1816</p> <p> <i>PANTHERA LEO SPELAEA</i> (GOLDFUSS, 1810)</p> <p>(FIGS 4–8)</p> <p> 1810 <i>Felis spelaea</i>. – Goldfuss: 277–280, table V, fig. 1.</p> <p> <i>Holotype:</i> <i>Felis spelaea</i> Goldfuss, 1823. Skull (Figs 4–8) of an adult male individual (Museum für Naturkunde der Humboldt-Universität Berlin No. MB.Ma.50948).</p> <p> <i>Paratype:</i> Lower jaw composed of two different individuals and of different sexes. At least one mandible clearly does not belong to the cranium because of its smaller size; the other has a strongly used canine, which indicates a very old individual, and would also not fit with the adult male skull (No. MB.Ma.50948).</p> <p> <i>Locus typicus:</i> Zoolithen Cave, Geilenreuth (cave No. D 106), Franconian Alb, Wiensent valley, Bavaria, southern Germany, Central Europe.</p> <p> <i>Stratigraphy:</i> ‘Upper layers above the main cave bear bone gravels’ after Goldfuss (1810, 1823). Weichselian/ Würmian, Upper Pleistocene (most probably OIS 3–6). OIS 3–6 after Rosendahl & Kempe (2004). Radiocarbon date on cave bear teeth 28.905 ± 255 BP (Poll, 1972).</p> <p> <i>Collection:</i> Museum für Naturkunde of the Humboldt- Universität Berlin.</p> <p> <i>Short description and discussion:</i> The nearly complete skull (Figs 5–8) and lower jaws cannot all be derived from one individual. The left mandible is about 3 cm shorter and more slender in its proportions. This half mandible would fit better with a lioness. The right mandible has a very heavily used canine, which again would not fit with the adult male skull. Several crania and even some lower jaws are present from this cave. The original skull has unique saggital crest damage. This is the result of a strong bite mark on the right parietal bone. This damage had undergone healing for a short time. The deep bite left a V-shaped notch in the centre of the crest. Only this skull of four possible candidates has these characteristics. Additionally, on the left side there is a modern hole damage in the skull, and the lower left jaw is missing most of the ramus, which must have occurred during the excavations. All these characteristics can be found only in one skull in the collection of the Museum für Naturkunde Berlin. The skull is 40.2 cm in total length and fits the description of skulls of male Late Pleistocene lions, which had skulls about 10 cm longer than those of female lions (cf. Altuna, 1981; Argant, 1988; Gross, 1992). The dentition was shown to be more complete in Goldfuss (1810). The lower jaw is now missing the right P 3 and all incisors have been lost. Similar losses can be observed in the upper incisors. Only the right I 2 now remains. It appears that the nasal bones have lost some anterior parts. The skull probably sustained modern frontal damage during which it unfortunately lost many teeth and other parts. Additionally, some parts in the symphyseal region of the mandible have been filled with plaster, especially around the canines.</p> <p> <i>Palaeopathological observations:</i> The skull of the male lion from Zoolithen Cave obviously had been severely bitten. The saggital crest was punctured up to 2 cm depth by the bite of another carnivore, and a large piece of bone must have been removed. Thus, the characteristic V-shaped notch in the middle of the saggital crest (Figs 8.2, 9A), which was clearly drawn by Goldfuss in 1810, can be clearly seen and is unique in all known lion skulls of the Zoolithen Cave. Additionally, a deep groove, 2 mm wide and about 5 cm long, starts from the V-shaped depression. This braincase damage must have been caused by the incisor of an attacking large carnivore. The bite was deep, and therefore it is possible that parts of the brain were damaged. The motor cortex (area motoria) is situated in this area (Fig. 9). The deep cut was in an early stage of the healing process with some callus production, but the animal cannot have survived for more than a couple of days. Even if the area motoria was not directly damaged, the callus healing process would have had an influence on the behaviour of the animal. The injured lion was possibly not able to move properly, having lost some of its motor capacity, but detailed palaeopathological studies, possibly with tomography, will need to be made to understand the exact injury and its impact.</p>Published as part of <i>Diedrich, Cajus G., 2008, The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo-ecological interpretation, pp. 822-831 in Zoological Journal of the Linnean Society 154 (4)</i> on pages 825-827, DOI: 10.1111/j.1096-3642.2008.00425.x, <a href="http://zenodo.org/record/5445628">http://zenodo.org/record/5445628</a&gt

The vertebrates from the Lower Ladinian (Middle Triassic)bonebed of Lamerden (Germany) as palaeoenvironmentindicators in the Germanic Basin

A marine/limnic vertebrate fauna is describedfrom the enodis/posseckeri Bonebed mixed in a bivalveshell-rich bioclastic carbonate rudstone at the easterncoastal margin of the Rhenish Massif mainland at Lamerden(Germany) within the western Germanic Basin (CentralEurope). The condensation layer is of Fassanian (Ladinian,Middle Triassic) in age. The vertebrate biodiversityincludes five different shark, and several actinopterygianfish species represented by teeth and scales. Abundantisolated bones from a small- and a large-sized pachypleurosaurNeusticosaurus species, which can be composedas incomplete skeletons, originate from dense populationsof different individual age stages. Important faciesindicator reptiles are from the thalattosaur Blezingeriaichthyospondyla which postcranial skeleton is reconstructedhypothetically using additional postcranialbones from similar aged various German localities. Thevertebrate biodiversity of the enodis/posseckeri bonebedof Lamerden reflect a limnic/fluvial freshwater influencedfauna (amphibians/terrestrial and marine reptiles) withdominance of normal saline marine influences. Macroalgaemeadow adapted placodont reptiles are absent inLamerden, as well as open marine-adapted ichthyosaurs,supporting a lagoon with fresh water influence position atthe Rhenish Massif mainland coast. In those contemporanousbrackish lagoons, which seem to be isochronousto northern Tethys lagoons of the Kalschieferzone at theMonte San Giorgio (Switzerland/Italy), small pachypleurosaurswere abundant prey in both regions for reptilepredators, especially large paraxial swimming alligatorhabitus-like Paranothosaurus, which even contain stomachcontents of pachypleurosaurs

Pleistocene Panthera leo spelaea (goldfuss 1810) remains from the Balve cave (NW Germany) – a cave bear, hyena den and middle palaeolithic human cave - and review of the Sauerland Karst lion cave sites

Pleistocene remains of Panthera leo spelaea (Goldfuss 1810) from Balve Cave (Sauerland Karst, NW-Germany), one of the most famous Middle Palaeolithic Neandertalian cave sites in Europe, and also a hyena and cave bear den, belong to the most important felid sites of the Sauerland Karst. The stratigraphy, macrofaunal assemblages and Palaeolithic stone artefacts range from the final Saalian (late Middle Pleistocene, Acheuléen) over the Middle Palaeolithic (Micoquien/Moustérien), and to the final Palaeolithic (Magdalénien) of the Weichselian (Upper Pleistocene). Most lion bones from Balve Cave can be identified as early to middle Upper Pleistocene in age. From this cave, a relatively large amount of hyena remains, and many chewed, and punctured herbivorous and carnivorous bones, especially those of woolly rhinoceros, indicate periodic den use of Crocuta crocuta spelaea. In addition to those of the Balve Cave, nearly all lion remains in the Sauerland Karst caves were found in hyena den bone assemblages, except those described here material from the Keppler Cave cave bear den. Late Pleistocene spotted hyenas imported most probably Panthera leo spelaea body parts, or scavenged on lion carcasses in caves, a suggestion which is supported by comparisons with other cave sites in the Sauerland Karst. The complex taphonomic situation of lion remains in hyena den bone assemblages and cave bear dens seem to have resulted from antagonistic hyena-lion conflicts and cave bear hunting by lions in caves, in which all cases lions may sometimes have been killed and finally consumed by hyenas. The lion remains, and not only in the Balve Cave seem to have been selected, as suggested by cranial and distal limb bone overrepresentations, which consist of 99 % of grown ups and with 58 % remains of dominantl females. Such limbs, and especially the pedal bone dominance, is not a criterion for “human hunt and fur import”, on the contrary, at hyena dens all prey remains are overrepresented by distal leg remains, a fact also very well known in the case of horse remains. The only articulated lion cub skeleton remain in the Sauerland Karst from the Wilhelms Cave might indicate a hyena kill that seems to be imported into the much frequented cub raising hyena den site

The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany : specialised cave bear predators of Europe

Remains of 13 individuals with 3/1 male/female ratio of the extinct Upper Pleistocene lion Panthera leo spelaea (Goldfuss, 1810) from the Zoolithen Cave near Burggeilenreuth (Bavaria, Germany) include the holotype skull and all paratype material. The highest mortality rate for the Zoolithen Cave lions is in their reproductive adult ages. Bite marks on lion bones or skulls are results of hyena activities, or rare cannibalism of lions under stress situations. Lions were possibly also killed in battles with cave bears during predation on hibernating bears in winter times. This cave bear hunt specialisation in caves overlaps with the ecological behaviour of cave bear feeding by Ice Age-spotted hyenas. Both largest Ice Age predators, lions and hyenas, had to specialise on feeding herbivorous cave bears in boreal forest mountainous cave rich regions, where the mammoth steppe megafauna prey was absent. This cave bear hunt by felids, and scavenging by hyenas and other large carnivores such as leopards and wolves explains why cave bears hibernated deep in to the European caves, for protection reasons against predators. Within such lion–cave bear and even lion–hyena conflicts in the caves lions must have been killed sometimes, explaining mainly the skeleton occurrences in different European caves

The Crocuta crocuta spelaea (Goldfuss 1823) population from the early Late Pleistocene hyena open air prey deposit site Biedensteg (Bad Wildungen, Hess, NW Germany), a contribution to their phylogenetic position, coprolites and prey

Four skulls, three lower jaws, a few incomplete postcranial bones and many coprolites of the early Late Pleistocene (Early Weichselian, 90,000 – 65,000 BP) ice age spotted hyena open air prey deposit site Biedensteg at Bad Wildungen (Hessia, NW-Germany) all show crack-, bite- and nibbling-marks as a result of cannibalism. Originally, the bones belong to three young adult to adult individuals. For the first time in Europe, a skull and postcranial bones belonging to a young animal of C. c. spelaea (Goldfuss 1823) of only a few days or weeks old are described and figured. The animal was possibly killed and for sure eaten by members of the same species. The material has been compared to remains from a younger Late Weichselian hyena population of Perick Caves (Sauerland, NW Germany). The hyenas from Biedensteg possess an upper first molar, in contrast to those from Perick Caves in which these are often lacking. At the Bad Wildungen-Biedensteg open air prey deposit site, the hyenas represent 6% of the mammoth steppe fauna bones. The site indicates a mixed diet consisting of all larger ice age mammals. The very high percentage (47%) of Coelodonta prey remains results of one disarticulated female and one young animal carcass, on which the hyenas fed strongly. Additionally, woolly rhinoceros bones indicate a specialization of the hyenas to large rhinoceros prey. This can be observed at other places, too, such as the hyena Perick Cave den. The abundant coprolites of the Biedensteg hyena population show different shapes, although most are droplet shaped or partly connected pellets which seemed to be of dry origin. Other coprolites, up to ten cm large, must have resulted from more soft dung. Many coprolites contain up to 1,5 cm small and well rounded bone compacta fragments, but also quite often bone spongiosa. The latter corresponds with the hyenas feeding on long bones of the woolly rhinoceros and steppe bison, which constitutes therefore another proof, next to the many chewing and gnawing marks at all prey bones. During the early Late Pleistocene, Biedensteg was a well frequented hyena open air den and prey deposit site close to the margin of a large sinkhole, filled with a shallow lake or muddy area, close to the small Wilde river. Also owls left many pellets at Biedensteg with thousands remains of micromammals, frogs and fishes

‘Neanderthal bone flutes’ : simply products of Ice Age spotted hyena scavenging activities on cave bear cubs in European cave bear dens

Punctured extinct cave bear femora were misidentified in southeastern Europe (Hungary/Slovenia) as ‘Palaeolithic bone flutes’ and the ‘oldest Neanderthal instruments’. These are not instruments, nor human made, but products of the most important cave bear scavengers of Europe, hyenas. Late Middle to Late Pleistocene (Mousterian to Gravettian) Ice Age spotted hyenas of Europe occupied mainly cave entrances as dens (communal/cub raising den types), but went deeper for scavenging into cave bear dens, or used in a few cases branches/diagonal shafts (i.e. prey storage den type). In most of those dens, about 20% of adult to 80% of bear cub remains have large carnivore damage. Hyenas left bones in repeating similar tooth mark and crush damage stages, demonstrating a butchering/bone cracking strategy. The femora of subadult cave bears are intermediate in damage patterns, compared to the adult ones, which were fully crushed to pieces. Hyenas produced round–oval puncture marks in cub femora only by the bone-crushing premolar teeth of both upper and lower jaw. The punctures/tooth impact marks are often present on both sides of the shaft of cave bear cub femora and are simply a result of non-breakage of the slightly calcified shaft compacta. All stages of femur puncturing to crushing are demonstrated herein, especially on a large cave bear population from a German cave bear den

Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears

The world’s first mounted skeletons of the Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) from the Sloup Cave hyena and cave bear den in the Moravian Karst (Czech Republic, central Europe) are compilations that have used bones from several different individuals. These skeletons are described and compared with the most complete known skeleton in Europe from a single individual, a lioness skeleton from the hyena den site at the Srbsko Chlum-Komín Cave in the Bohemian Karst (Czech Republic). Pathological features such as rib fractures and brain-case damage in these specimens, and also in other skulls from the Zoolithen Cave (Germany) that were used for comparison, are indicative of intraspecific fights, fights with Ice Age spotted hyenas, and possibly also of fights with cave bears. In contrast, other skulls from the Perick and Zoolithen caves in Germany and the Ur?ilor Cave in Romania exhibit post mortem damage in the form of bites and fractures probably caused either by hyena scavenging or by lion cannibalism. In the Srbsko Chlum-Komín Cave a young and brain-damaged lioness appears to have died (or possibly been killed by hyenas) within the hyena prey-storage den. In the cave bear dominated bone-rich Sloup and Zoolithen caves of central Europe it appears that lions may have actively hunted cave bears, mainly during their hibernation. Bears may have occasionally injured or even killed predating lions, but in contrast to hyenas, the bears were herbivorous and so did not feed on the lion carcasses. The articulated lion skeletons found in cave bear dens deep within caves scattered across Europe (such as those from the Sloup, Zoolithen and Ur?ilor caves) can therefore now be explained as being the result of lions being killed during predation on cave bears, either by the cave bears defending themselves or as a result of interspecific fights

Hermann’s Cave (Germany) – A Late Pleistocene Cave Bear Den

The northern German Hermann’s Cave below the 1.141 a.s.l meters high Brocken peak in the centre of the Harz Mountains (Saxony-Anhalt) was discovered by chance during road construction work at the village Rübeland in the year 1866. It was explored and a first map was presented with first cave descriptions 1889 by the geologists Prof. Dr. J.H. Kloos and Prof. Dr. M. Müller from the Brunswick University. In 1890 the cave became one of the historically opened European show caves and is one of the largest tourist caves in Germany counting about 75.000 visitors per year. Palaeontological and archaeological pioneering research was made by the biologist Prof. Dr. W. Blasius from the Brunswick Natural History Museum, who was active from 1892 to 1901 with opening a small museum in front of the cave. The Hermann’s Cave bear den belongs to one of the three bone-rich important and most northern European cave bear den sites being situated opposite the Baumann’s Cave and not far from the Unicorn Cave in the Harz Mountain. In contrast to the other two mentioned caves, which were used by cave bears and carnivores such as Neanderthals in the Middle Pleistocene, the Hermann’s Cave was accessible for cave bears, carnivores and even humans due to the starting pre-LGM glaciations of the Brocken peak and cave entrance collapse processes only in the Late Pleistocene. At this time, Ice Age steppe lions and Late Palaeolithic Cromagnon humans hunted different cave bear species deep in the cave