Halle (Saale), Germany


Halle (Saale), Germany
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The classical descriptions of Middle Triassic marine Placodus gigas Agassiz, 1833 (Reptilia) from the Germanic Basin of central Europe as being shell-crushing durophagous placodontids are revised in this paper through analyses in convergent anatomy. In particular, the jaw morphologies of three placodontid genera, Paraplacodus, Placodus and Cyamodus, are compared to those of dugongs (Mammalia) such as the central European Tertiary Halitherium schinzii and the modern Dugong dugon of the Arabian Gulf. The anatomies of Paraplacodus, Placodus and Cyamodus exhibit convergences to Halitherium and Dugong. Whereas mammalian dugongs developed pachyostotic thoracic ribs to enhance their body weight, the placodontid reptiles achieved a similar result in different ways: Paraplacodus developed enlarged thoracic ribs; Placodus had pachyostotic gastral ribs, and Cyamodus had a thoracic osteoderm shield. The teeth of the placodontids are also convergent with those of Halitherium and Dugong in their general function and jaw morphology. Whereas Halitherium and the modern Dugong possess a horny oral pad and counterpart, and a specialized rasp-like tongue with which to grind the seagrass and its roots, placodontids had large teeth that covered the whole of their upper and lower jaws forming a similar crushing or grinding pad. Both of the extinct groups must have fed on sea-plants, as does the modern Dugong, although Halitherium possibly fed on both seagrass and macroalgae. A study of the wear stages of many Placodus teeth, skulls and jaws has revealed a large proportion of highly worn anterior teeth, indicating a usage similar to that of the procumbent front teeth of modern Dugong which are used to scrape plant roots from the sea-floor. In contrast, highly worn (wear stage 3) teeth are rare (0.5%) amongst all other upper palatal, maxillary, or lower-jaw dentary teeth, suggesting a relatively soft diet. Placodus must have used their broadly spaced anterior teeth, to dig macroalgae from carbonate sands in shallow marine, sand bar environments; indirect evidence for the existence of such environments is provided by the benthic communities of the Germanic Basin and the northern Tethys. Sea-plants would have been only crushed and swallowed by the placodontids and not chewed with jaw rotation, in a similar feeding strategy to that used by modern Dugong feeding on seagrass. © 2009 Elsevier B.V. All rights reserved.

In the greenhouse world of the Early Upper Cretaceous of Europe giant cephalopod shells of the hemibenthic ammonite Puzosia formed huge accumulations. Their shells indicate adaptations in the body chamber morphology depending on the environment. On the sea floor, deposited ammonite shells scoured up to 50 cm depth which caught more and more large shells over extended periods of time. They built up to five square meter extended scour troughs in which the shells are enriched in chain-, fan- and fan layer orders with a maximum accumulation of 24 cephalopods. Between these hundreds of other macrofaunal remains accumulated. The ammonite shells were benthic islands and minibiotopes in carbonate soft- to firm ground environments along a submarine swell in the southern North Sea Basin of Central Europe. The empty shells encrusted by different epizoans sheltered and protected also crustaceans, which undermined the empty shells by bioturbation. Small and extremely rare squamate reptiles of the marine Dolichosaurus longicollis possibly took shelter or fed within these unique benthic submarine depression "islands".

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. © 2011 Taylor & Francis.

Remains from at least seven individuals of the Late Pleistocene Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) from the Teufelskammer Cave in the Neandertal valley (North Rhine-Westphalia, northwest Germany) are described. The small cave was a well-frequented hyena den of the Early to Middle Late Pleistocene which was only 100m from the famous small Feldhofer Cave, where the first Neandertal human skeleton was found. The high amount of hyena bone material (37%) and its strongly chewed and incomplete prey remains of the mixed mammoth steppe and boreal forest megafauna prove one more of 11 recently known hyena den caves in the Rhenish Massif. Hyenas and cave bears have used the cave, but Neandertal humans lived possibly not at the same time in the same valley. Although hyenas occupied mainly the smaller caves such as the Teufelskammer Cave, humans preferred large portal cave entrances such as in the Neandertal valley with the Small Feldhofer Cave. © 2011 Taylor & Francis.

European leopard sites in Europe demonstrate Early/Middle Pleistocene out of Africa lowland, and Late Pleistocene Asian alpine migrations being driven by climatic changes. Four different European Pleistocene subspecies are known. The final European Late Pleistocene "Ice Age leopard" Panthera pardus spelaea (Bächler, 1936) is validated taxonomically. The skull shows heavy signs of sexual dimorphism with closest cranial characters to the Caucasian Panthera pardus ciscaucasica (Persian leopard). Late Pleistocene leopards were distributed northernmost, up to S-England with the youngest stratigraphic records by skeletons and cave art in the MIS 2/3 (about 32,000-26,000BP). The oldest leopard painting left by Late Palaeolithics (Aurignacians/Gravettians) in the Chauvet Cave (S-France) allows the reconstruction of the Ice Age leopard fur spot pattern being close to the snow or Caucasian leopards. The last Ice Age glacial leopard habitat was the mountain/alpine boreal forest (not mammoth steppe lowland), where those hunted even larger prey such as alpine game (Ibex, Chamois). Into some lairs, those imported their prey by short-term cave dwelling (e.g. Baumann's Cave, Harz Mountains, Germany). Only Eurasian Ice Age leopards specialized, similar as other Late Pleistocene large felids (steppe lions), on cave bear predation/scavenging partly very deep in caves. In Vjetrenica Cave (Dinarid Mountains, Bosnia Herzegovina), four adult leopards (two males/two females) of the MIS 3 were found about two km deep from the entrance in a cave bear den, near to one cave bear skeleton, that remained articulated in its nest. Leopards died there, partly being trapped by raising water levels of an active ponor stream, but seem to have been killed possibly either, similar as for lions known, in battles with cave bears in several cave bear den sites of Europe (e.g. Baumann's Cave, Wildkirchli Cave, Vjetrenica Cave). At other large cave sites, with overlap of hyena, wolf and dhole dens at the cave entrances, leopard bones with bite damages indicate their remains to have been imported and consumed by predators in alpine regions due to reduced prey availability. The best models for the competition/taphonomy of large predators - felids, hyenids, canids - within large cave bear dens of Europe is represented in combination of the Zoolithen Cave and Vjetrenica Cave taphonomy. © 2013 Elsevier Ltd.

The upper Pleistocene Ice Age spotted hyena remains (number of bones per species (NISP) = 206; minimum individual number (MNI) = 7 young, 12 adult) from the German Zoolithen Cave include the Crocuta crocuta spelaea (Goldfuss 1823) holotype skull, and all cranial and postcranial paratypes which were found in the 'cave bear bonebeds'. Those bones are on secondary positions in the vertical shafts especially of the central cave part, where they were redeposited due to quick flooding events of the Wiesent River at the final late Pleistocene. The young animal bones (NISP = 13%) indicate a natal/birth den site. Cannibalism within the hyenas is demonstrated by several skulls and few long bones. The scarcity of steppe megafauna hyena prey (NISP = 1%, woolly rhinoceros remains) and high amounts of damaged cave bear long bones (17%), similarly preserved at many other dens in Central Europe, indicate prey specialisation of hyenas (and other carnivores such as steppe lions, leopards and wolves) onto cave bear feeding in medium-high mountainous regions in the caves. This is the result of the rare mammoth steppe megafauna in such late Pleistocene boreal forest and cave-rich environments, which could have been hunted by the Zoolithen Cave hyenas only during migrations along the Wiesent River valley. © 2011 Copyright Taylor and Francis Group, LLC.

Diedrich C.G.,PaleoLogic
Biological Journal of the Linnean Society | Year: 2011

A systematically excavated track site in a 243.5Myr old Middle Triassic (Karlstadt Formation, Pelsonian, middle Anisian) intertidal carbonate mud-flat palaeoenvironment at Bernburg (Saxony-Anhalt, central Germany) has revealed extensive horseshoe crab trackways attributable to the KouphichniumNopsca, 1923 ichnogenus. The exposed track bed of a Germanic Basin-wide spanned intertidal megatrack site is a mud-cracked biolaminate surface on which detailed tracks have been preserved because of rapid drying and cementation as a result of high temperatures, followed by rapid covering with a protective layer of arenitic storm or tsunami sediments. The different trackway types and their orientations have allowed a tidal sequence to be reconstructed, with the initial appearance of swimming horseshoe crabs followed by half-swimming/half-hopping limulids under the shallowest water conditions. The Bernburg trackways, which have mapped lengths of up to 40m, were all produced by adult animals and exhibit a variety of shapes and patterns that reflect a range of subaquatic locomotion behaviour more typical of mating than of feeding activities. The closest match to the proportions and dimensions of the horseshoe crab tracks at Bernburg is provided by the largest known Middle Triassic limulid Tachypleus gadeai, which is known from the north-western Tethys in Spain. The horseshoe crab body fossils recognized in the German Mesozoic intertidal zones, instead, are from juveniles. The uniformly adult size indicated by the trackways therefore suggests that they may record the oldest intertidal reproductive zones of horseshoe crabs known from anywhere in the world, with the track-makers having possibly migrated thousands of kilometres from shallow marine areas of the north-western Tethys to reproduce in the intertidal palaeoenvironments of the Germanic Basin. Chirotherium trackways of large thecodont archosaurs also appeared on these flats where they appear to have fed on the limulids. With the tidal ebb, smaller reptiles such as Macrocnemus (Rhynchosauroides trackways) appeared on the dry intertidal flats, probably feeding on marine organisms and possibly also on horseshoe crab eggs. © 2011 The Linnean Society of London.

The remains of a large population of Late Pleistocene Ice Age spotted hyenas (Crocuta crocuta spelaea Goldfuss 1823) are described from the Rösenbeck Cave in the Sauerland Karst of Germany. They include four skulls and 79 other skeletal parts, mainly from adult to senile animals, making this an important Late Pleistocene hyena cave-den site in Europe. The skulls have been compared with 30 other hyena skull specimens from open air and cave-den sites in central Europe (Germany, Austria, the Czech Republic and Romania) in order to achieve an understanding of sexual dimorphism in the crania of Ice Age spotted hyenas from the Upper Pleistocene cold period (Weichselian/Wuermian), and the types of injuries that they acquired during their lifetimes. Three different types of cranial shape have been distinguished, one of which appears to have been a consequence of pathologies that developed in response to injuries caused by bites received during the animal's lifetime, as a result of either intraspecies fights or fights with lions. Although cave bears penetrated to great depths within the Rösenbeck Cave for hibernation purposes, hyenas appear to have utilized only a short section of the cave that branched off directly from the entrance area. Hyena cub material is scarce, suggesting that this area was used as a communal den rather than for cub rearing. Bones exhibiting gnaw marks, representing prey imported by hyenas, are also rare but include horse (Equus caballus przewalskii) and reindeer (Rangifer tarandus) remains. The scarcity of bones from hyena prey suggests that this cave was not used as a food storage site. Some Ursus spelaeus cave bear remains, including skulls, show evidence of having been gnawed, chewed and cracked by hyenas, indicating that the hyenas periodically fed on cave bear carcasses in a specialization response to the mammoth steppe megafauna absence of the boreal mountain forest regions. © 2011 The Linnean Society of London.

The phylostratigraphy, taphonomy and palaeoecology of the Late Cretaceous neoselachian Ptychodus of northern Germany appears to be facies related. Ptychodus is not present in lower Cenomanian shark-tooth-rich rocks. First P. oweni records seem to relate to middle Cenomanian strata. P. decurrens appears in the middle to upper Cenomanian mainly in non-coastal environments of the shallow marine carbonate ramp and swell facies which isolated teeth were found partly in giant ammonite scour troughs on the Northwestphalian-Lippe High submarine swell in the southern Pre-North Sea Basin. They are recorded rare in deeper basin black shales facies (upwelling influenced, OAE Event II). P. polygyrus seems to be restricted to upwelling influenced basin and deeper ramp facies mainly of the uppermost Cenomanian and basal lower Turonian (OAE II Event). P. mammillaris is mostly represented during the lower to middle Turonian in the inoceramid-rich ramp and the near shore greensand facies along the Münsterland Cretaceous Basin coast north of the Rhenish Massif mainland. Finally, P. latissimus is recorded by two new tooth sets and appears in the upper Turonian basin swell facies and the coastal greensands. Autochthonous post-Turonian Ptychodus remains are unrecorded in the Santonian-Campanian of Germany yet. Reworked material from Cenomanian/Turonian strata was found in early Santonian and middle Eocene shark-tooth-rich condensation beds. With the regression starting in the Coniacian, Ptychodus disappeared in at least the Münster Cretaceous Basin (NW-Germany), but remained present at least in North America in the Western Interior Seaway. The Cenomanian/Turonian Ptychodus species indicate a rapid neoselachian evolution within the marine transgression and global high stand. A correlation between inoceramid shell sizes, thicknesses and their increasing size during the Cenomanian and Turonian might explain the more robust and coarser ridged enamel surfaces in Ptychodus teeth, if Ptychodus is believed to have preyed on epifaunistic inoceramid bivalves. © 2012 Elsevier Ltd.

A Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) population from the cave bear den Sloup Cave, Moravia (Czech Republic) consists of mainly adult/senior and few cub/juvenile remains and coprolites, and 139 prey bones. Hyenas used the Nicová Cave branch that is connected to the entrance area mainly as a communal den site. Prey bone damage is most visible on the imported woolly rhinoceros remains. The partly excavated prey bone accumulation consists of a single woolly mammoth Mammuthus primigenius (Blumenbach 1799) tooth (2%), mainly Coelodonta antiquitatis (Blumenbach 1807) remains (16%), 4% Bos primigenius (Bojanus 1827) and 1% each of Megaloceros giganteus (Blumenbach 1799) and Rangifer tarandus (Linnaeus 1758). The other carnivores such as Panthera leo spelaea (Goldfuss 1810), Gulo gulo (Linnaeus 1758) and Canis lupus (Linnaeus 1758) subsp. are less represented (1-3%). Wolverines might have been imported also as prey remains, whereas wolves also possibly used this cave on a short-term basis, whereas steppe lions seem to have preyed upon cave bears deeper in the cave periodically, where even skeletons of P. leo spelaea were found in the Elisabeth Cave part. © 2012 Copyright Taylor and Francis Group, LLC.

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