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In Germany tens to hundreds of kilometres-laterally distributed sedimentological heterogene crinoid accumulation bed with partly articulated Encrinus liliiformis specimens is dated within a complete trans-regression cycle of higher order into the upper Illyrian, Anisian (Middle Triassic, pulcher biozone). The crinoid bearing layer differs much from under- and overlaying tempestite parasequence sets of the Meisner Formation. The tsunamite layer consists of different laterally grain size sorted facies types ranging from unsorted bioclastic floatstones of large-scaled antidune megaripples, which continue in oscillation-ripple-marked oolithic grainstone surfaces that end into crinoid filled scour troughs in wacke/mudstones (=horizontal grain sorting in backwash direction). The tsunamite bed of Lamerden that consists of mixed reworked oolithic bar to crinoid ramp facies areas (sediments/biota to the outer ramp facies) can be correlated to a crinoid bed of Bissendorf within northwestern Germany at minimum (about 50 km) and appears within the tectonical deepening period of the Central European intracratonic Germanic Basin. The heterotope crinoid taphocoenosis was built in the outer ramp Tonplatten facies, in the deepest part of the Germanic Basin (Hesse-Depression) due to backwash. The mixed macro fauna originate from the crinoid bioherms and terebratulid hardgrounds of the inner/middle ramp facies (such as the sediments). From 350 analysed crinoid individuals (of 1500), only 1 % are complete with 120–160 cm short-stemmed ecotypes of the inner ramp crinoid meadows which settled along the sand bars basinwards. The Lamerden crinoids are scattered or are concentrated differently condensed in scour trough-like structures on wackestones/mustone areas and more rare or scattered on osciallation-ripple marked grainstones and allow only local bottom current estimations. In interfingering floatstones, more rare crowns are found unsorted within dominat chaotic oriented terebratulids. Most crinoid specimens (83 %) were found with relic stems and the others (16 %), only as crowns. The echinoderms survived a tsunami event, for at least a few hours/days under 10–30 cm of mud coverage. Only 1 % of the crowns have opened arms (=postmortal embedding); all others (99 % are closed crowns) are “buried alive” specimens. The crinoids closed the nutrition canal immediately and started regeneration of the distal stem for new attachment purposes; therefore, those have all rounded distal stems. The northern German Lamerden crinoid layer is most similar to the southern German Neckarwestheim site, but is different in its origin to other German sites, where crinoids have been found around bioherms or with long-stemmed specimens not far from their patch-reef like habitats in channels of the upper middle ramp facies. The tsunamite demonstrate the problem of the “Triassic time scale” which is tempestite and Milankovitch cycle based on an intracratonic basin, because there seem to be several further tsunamite bioclastic layers, which were misidentified as “tempestites”.

Four new saber-tooth cat ( Homotherium ) sites in Germany with new dental and postcranial bone material are different in their taphonomic context: 1. The Archaeological Middle Palaeolithic (MIS 9e-Interglacial) Schoningen Lake site with remains of a cub carcass, 2. The Middle Palaeolithic (MIS 5e-9) Archaeological/cave bear den site of Balve Cave yielding a lower canine tooth of an older individual, 3. The Zoolithen Cave (MIS 3–9) cave bear/hyena den with one distal half humerus of an adult, 4. The Ketsch open air Rhine River terrace site which has provided another distal humerus of an adult saber-tooth cat. Whereas only the Schoningen site is precisely dated as Holsteinian Interglacial (approx. 330.000–315.000 BP), all other material seems to come from the same Middle Pleistocene warm period, or few younger Saalian interstadials (MIS 7a, e) deposits, and did not extend over the last MIS 7 glacial into the Late Pleistocene. Homotherium as hyena-like slow moving cat seems to have disappeared within the Saalian due to competition with other scavengers like Ice Age spotted and brown hyenas ( Crocuta crocuta praespelaea/ultima and Pachycrocuta brunnea mosbachensis ). The juvenile saber-tooth cat cub from Schoningen might be in archaeological context or represent only a carnivore kill. At the Zoolithen Cave, the single bone must have been imported into a hyena prey bone assemblage. The situation is possibly similar at the two other sites Ketsch and Balve Cave. The formerly described Schoningen “saber-tooth cat” humerus is revised, such as other opposite as lion humeri described material from different European sites. The presence of the well-developed supracondylar ridge distinguishes Homotherium well from Middle/Late Pleistocene lions Panthera leo (e.g. spelaea, fossilis ). The Schoningen lion humrus has been chew-cut first most probably by a stripped hyena whose cutting scissor teeth produced a diagonal bite cut and P 4 /M 1 impact marks around the trochlea. 1–2 mm small, mostly triangular-oval bite marks on the lion humerus shaft compacta results from a second scavenger and not from “Neanderthal tool use”. Those bite mark sizes are produced mainly of the upper molar teeth of a the red wolf Cuon alpinus subsp. (or small fox Vulpes praecorsac ), which were present in the region within the Holsteinian/Saalian.

A total of 366 bones are arranged to six incomplete individual skeletons (=MNI) of different age classes and taphonomic preservation from the Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) hyena den Srbsko Chlum–Komin Cave, Czech Republic, Bohemian Mountains (Central Europe). The hyena assemblage counts no siblings, 37% cub, 33% early adult/subadult and 30% adult/senile individuals. A senile scavenged individual demonstrates the cannibalism within the cave. About 10% of the 3569 megafauna bones are from hyenas which larger amounts are typical at communal/prey depot den sites. This site is different to three other larger natal/birth and communal or prey depot hyena den cave sites in Czech Republic. The Chlum–Komin Cave hyena prey bone assemblage (=NISP) consists, similar as at two other Czech Republic sites, mainly of Equus ferus przewalskii (50%), including pregnant females being hunted in spring/early summer. The fewer amounts of Coelodonta antiquitatis (4%), Bison priscus (2%), Rangifer tarandus (15%), and alpine Rupicapra rupicapra (2%) or Capra ibex (1%) and the absence of mammoth correlate to hyena den bone assemblages in middle high elevated mountain regions of northern/central Europe. The rareness of cave bears in the Bohemian Mountains, on which hyenas specialized to feed additionally in European mountain regions, explains predominant predation on horses.

Newly excavated Middle Triassic reptile Chirotherium, Rhynchosauroides and horseshoe crab Kouphichnium imprints from the uppermost Pelsonian megatracksite Bernburg, Germany (Central Europe) represent different behavioural trackways. The Chirotherium/Rhynchosauroides/Kouphichnium track assemblage of the lower intertidal carbonate flats is dominated by well-known horseshoe crab tracks of various subaquatic benthic movements which were all left on mud-cracked biolaminites in reproduction coastal zones. Most abundant reptile Rhynchosauroides trackways (50 % of reptile tracks) are oriented more beach-parallel (to Rhenish Massif Island) as a result of food searching along the intertidal seismic-influenced coast (16 slickensided biolaminate layers within 2 m biolaminates due to Alpine fold belt and Germanic intracratonic Basin tectonics), most probably for feeding on high abundant horseshoe crab eggs resulting from mating events in the lower intertidal. Large basal archosaur predators (?Ticinosuchus) produced large Chirotherium trackways (25 % of reptile tracks) which are represented in the new material with exceptional unusual trackways (slipping and side steps), but in directions to and from the beach to the Rhenish Massif main land, indicating predation on small reptiles. Finally, medium-sized Chirotherium trackways of subadult animals are present (25 % of reptile tracks). One trackway has a unique “feeding place” area, indicating the unexpected either predation, playing or feeding of a basal archosaur on a horseshoe crab. The horseshoe crab migrations from the North Tethys onto the extended mating intertidal mud flat beaches of the Germanic Basin, and recently known even into Chinese lagoons migrating limulid populations of different horseshoe crab species must have caused a global and seasonal food chain reaction, whereas Macrocnemus reptiles (Rhynchosauroides trackmaker) might have fed only on Millions of their eggs.

A remarkable Middle Triassic (Pelsonian, Anisian) fossil track site at Bernburg in Central Germany permits reconstruction of a tetrapod fauna at the point of recovery when early poposaur quadruped dinosaurs diversificated globally. The site has yielded numerous tetrapod and thousands of arthropod horseshoecrab tracks from several levels in intertidal biolaminites with intercalated storm and seismic-influenced carbonates. The tetrapod tracks, Procolophonichnium, Rhynchosauroides, Chirotherium, and Isochirotherium, are assigned to smaller Archosauromorpha and large Archosauria. The largest chirotheroid tracks Isochirotherium herculis (Egerton in Proc Geol Soc London 3:14–15, 1838) reach up to 350-mm-long pes sizes and 120-mm-long manus imprints. Those were most probably made by a 5-m-long poposauroid archosaur like Arizonasaurus, based on matching skeleton anatomy and trackways including a new three-dimensional model of those giants in locomotion. A first German poposauroid bone record is added herein with a scapula from shallow marine carbonates of similar Pelsonian ages. These crurotarsan archosaurs were at the top of their food chains globally, and fed most probably on smaller tetrapods (Hescheleria, Macrocnemus), the latter also represented well by long behavioural escaping Rhynchosauroides trackways. The smaller reptiles may have fed especially on “Millions” of horseshoecrab eggs in those arthropod reproduction seasons in the intertidal carbonate mud flats. Such tidal flats and sabkhas extended over hundreds of square kilometres surrounding the Germanic Basin in Central Europe in the Middle Triassic, representing a drastic change in the palaeoenvironment of central Pangaea, and a possible trigger for continuing diversification of archosauromorphs and eventually dinosaurs. The horseshoecrab reproduction seasons seem to have caused a food chain reaction along the southern Pangaean (northern Tethys) coast in carbonate mud flats between Northern America, Africa, Europe and China, with possibly long-distance migrations of poposauroids such as Arizonasaurus, whose bones were found in terrestrial to shallow marine carbonate deposits in terrest to coastal context.

An Isurus denticulatus (Glickman, 1957) shark skeleton from the late Turonian (Late Cretaceous) of Germany is described within a diverse upwelling influenced fish fauna of northern Germany, Europe. It was found in the turbiditic marly limestones at the submarine Northwestphalian Lippe Swell in the southern Proto-North Sea Basin. Compared to modern mackerel shark Isurus oxyrinchus Rafinesque, 1809, including cranial denticles, this allows a revision of the younger synonym “Cretoxyrhina Glickman, 1964”. Within the Cretaceous Isurus, a loss of the lateral tooth cusps and nutritive clefts of the roots (considered as plesiomorphic character of the Lamnidae) took place from the Albian (Early Cretaceous) to the Campanian (Late Cretaceous). The tooth morphology changed during the Albian-Turonian from a tearing (I. denticulatus) to a cutting (I. mantelli) type (Coniacian-Campanian). The complete lateral cusplet and symphyseal teeth reduction in Isurus at the end of the Cretaceous seem to be a result of the coevolutionary changing feeding habits of a worldwide expanding shark. In a second evolutionary adaptation, parallel to the new radiation of marine mammals (Paleocene/early Eocene), from Isurus, the white shark ancestors (Carcharodon) seem to originate. In another radiation from Isurus, coevolving with appearance of dolphins and further marine mammal evolution within the Middle Miocene, a second time Isurus developed serrated teeth (I. escheri).

Late Pleistocene spotted hyenaCrocuta crocuta spelaea(Goldfuss, 1823) and steppe lionPanthera leo spelaea(Goldfuss, 1810) were top predators in Central Europe. The fossil record (2.303 hyena/1.373 lion bones = ratio 3/1) from 106 cave and open air sites demonstrates comparable associations to modern African hyenas/lions resulting in competition about prey and territory. Cannibalism within extinct spotted hyenas is well documented, including two individual skeletons. Those hyenas produced bone accumulations at dens. Feeding specializations on different megamammal groups are demonstrated for Late Pleistocene hyenas whose prey partly overlaps (e.g., cave bears) with those of lions and wolves. At most fossil sites, 1–3% of the lion remains indicate scavenging of lions by hyenas. The larger Late Pleistocene felids focussed on cervids (reindeers specialization during the high glacial = LGM), on bovids (steppe bison/aurochs), and possibly on saiga antelope and on the cave bear, hunting deep in caves during their hibernations and targeting cubs. The cave bear feeding was the target of all three top predators (lions, hyenas, and wolves) in the Late Pleistocene boreal forests which caused deathly conflicts in caves between them, especially with lions/hyenas and herbivorous cave bears that have no modern analogue.

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 ( Bachler, 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 32,000–26,000 BP). 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.

The scavenging activities of hyenas and lions at the Neumark-Nord Lake 1 (Central Germany) site on straight tusk elephant carcasses is commonly documented by slight bone damages and initial skeleton partitioning. Canine bite marks are found at several joints of elephant long bones, as well as on the vertebrae and skulls. Deep bite and scratch marks from 5 to 7 mm in width made by large carnivore canine teeth (Panthera leo spelaea, Crocuta crocuta spelaea) on the ventral vertebral columns and pelvises of two Palaeoloxodon antiquus straight-tusked elephant skeletons suggest that the intestines and inner organs were consumed by both large lion/hyena predators as is commonly observed when modern African lions and spotted hyenas feed on contemporary elephant carcasses. A scavenging strategy can be demonstrated using 24 partly preserved Palaeoloxodon skeletons, where Ice Age spotted hyenas appear to be the main scavenger (95 %) and sole skeleton decompositor such as bone destructor. While Crocuta may feed anywhere on an elephant carcass, they are known to specialize to eat the complete feet, whereas scavenging from the anus into the body cavity (intestine/internal organ feeding) is more typical of lions. Shallow water at Neumark-Nord Lake 1 prevented hyenas from completely scavenging elephants; thus, bone damage at this site was constrained to initial stages of destruction and final carcass partitioning by high water stands and floods.

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 Munsterland 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 Munster 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.

The early white shark Carcharodon Smith, 1838 with the fossil Carcharodon auriculatus (Blainville, 1818) and the extinct megatooth shark Otodus Agassiz, 1843 with species Otodus sokolovi (Jaeckel, 1895) were both present in the European proto North Sea Basin about 47.8 - 41.3 m.y. ago (Lutetian, early Middle Eocene), as well as in the Tethys realm around the Afican-Eurasian shallow marine habitats. Both top predators developed to be polyphyletic, with possible two different lamnid shark ancestors within the Early Paleocene to Early Eocene timespan with Carcharodon (white shark line-age) and Otodus (megatooth shark lineage). Their sawblade teeth developed during the early Paleogene as the result of adaptation to feeding on various marine new rising mammals, coinciding with three main waves of evolutionary emergence of seals, sirenians, and whales in parallel with the evolution of these large predatory sharks. Megatooth sharks specialized in hunting whales and sirenians only on the coastal shelves of warm oceans and disappeared globally in the Pleistocene due to climate change and ocean cooling. The cold-water adapted early white sharks have survived until the present day with body temperate change adaptation in warm to temperate oceans and are proposed to have specialized on coastal seal hunting already50 m.y. ago.

Newly discovered remains of the early Middle Eocene (Lutetian) sirenian Protosiren (Protosirenidae) in shark tooth rich conglomerates from a coastal delta environment northwest of the European Rhenish Massif at Furstenau (northwestern Germany), represent the most northerly occurrence of this genus whose global distribution was generally restricted to warm waters. Its presence of the remains so far north can be explained by seasonal inflow of warm Tethys surface water into the cool, upwelling-influenced, basin. The existence of two discrete centers of sirenian evolution can be explained by the opening of the Atlantic and the upwelling that separated the North American warm water faunal province from those of Africa and Eurasia. A slightly modified evolutionary model is presented in which the oldest Early Eocene manatee sirenians evolved in the Caribbean of Central America. Protosiren, however, appears to have developed polyphyletically along the African coastline of the Tethys, and represents the oldest known dugong ancestor. Younger (Oligocene) European sirenian skeletons of Halitherium and Anomotherium are included in the phylostratigraphic model in which sirenians had generally reduced their teeth by 28 Ma as an adaptation for feeding on sea-plants (macroalgae/seagrass). Teeth from early megatooth sharks, which preyed on sirenians, have been recorded from shallow marine Eocene and Oligocene coastlines of the southern proto-North Sea Basin, and shark bite marks have been found on sirenian skeletons.

Crocuta crocuta spelaea ( Goldfuss, 1823 ) remains (NISP = 50) are present in the early to middle Upper Pleistocene Emscher River terrace open air den site along the Rhine–Herne Canal near Bottrop (Westphalia, NW Germany). The population includes bones from cubs and bones with pathological features from old animals but is predominantly made up of adult hyenas (NISP = 3820) found within the glacial mammoth steppe fauna of the Munsterland Bay Lowlands. A larger quantity of woolly rhinoceros ( Coelodonta antiquitatis ) bones (NISP = 1601) are present than in other bone-rich open air hyena den sites, of which about 67% are damaged. This damage can be shown to have resulted mainly from hyena activities, and shows repetitions of the same three stages of destruction on the massive woolly rhinoceros bones. Those bone shafts that the hyenas were unable to break were left with irregular jagged margins. The taphonomy of bones from the Bottrop open air site indicates that the rhinoceros body parts had only been transported over short distances, in contrast to those from the Perick Caves hyena den where longer transport distances resulted in a higher proportion of limbs compared to other body parts. The large woolly rhinoceros was an important second megafauna prey for hyenas in the lowland areas (after the woolly mammoth), in contrast to the nearby Sauerland Karst mountain areas that are rich in hyena dens and their associated bone assemblages, although the higher proportions of rhinoceros bones present is also to some extent a result of the incomplete destruction of their massive bones. The proportions of woolly mammoth and woolly rhinoceros remains in the hyena prey bone assemblages gradually decrease from the lowlands towards the nearby Sauerland Karst mountain regions. In these mountainous boreal forest regions cave bears were instead the main food source for hyenas, as result of the scarcity or absence of mammoth steppe fauna. The large bone accumulations along the Emscher River terraces near Bottrop can mostly be attributed to the activities of hyenas, which were responsible for the repeated incomplete state of bone preservation. The bone accumulations are predominantly of leg bones and include a relatively high proportion of hyena bones which, together with the faunal composition and the size of the accumulations, indicate a mixed long-term use of the terraces along the Emscher River as an extensive communal hyena den and prey storage site, and also probably somewhere in the area, as a birth den. Some of the bone material may also have been accumulated by natural processes, or even by Neanderthals.

Chirotherid footprints of Synaptichnium, Chirotherium and Isochirotherium appeared during the late Early (Aegean) to early Late (Carnian) Triassic in central Europe. These taxa are partly revised herein, using both perfect and variably preserved tracks, and very long trackways from an upper Pelsonian intertidal-flat megatracksite of the Germanic Basin coast Pelsonian (Karlstadt Formation). The global Middle Triassic distribution of those footprints suggests seasonal migrations across Pangaea of possible archosauriform reptile trackmakers, such as Euparkeria, Ticinosuchus, Arizonasaurus and Batrachotomus, caused by horseshoe-crab mass migrations into tidal-flat beach reproductive zones in the Germanic Basin. Such seasonal migrations may even suggest a Pangaea-wide food-chain reaction, possibly including the mobilization of fish, marine and terrestrial reptiles, and of which situation the Germanic Basin intertidal-flats is a globally unique example.

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 Nicova 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-...

The Middle Triassic fossil reptile localities near Bayreuth (Bavaria, southern Germany) consist of shallow marine autochthonous glauconitic marls and terebratulid-rich tempestite carbonates of the newly defined Bindlach and Hegnabrunn formations. Single bones and incomplete skeletons of marine reptiles have been recorded in bone beds within in the Illyrian and Fassanian stages. These include the remains of the sauropterygians Neusticosaurus sp., Lariosaurus cf. buzzii [1], Nothosaurus mirabilis [2], Paranothosaurus giganteus [2], Placodus gigas [3], Cyamodus rostratus [4], Cyamodus münsteri [5], Pistosaurus longaevus [6], and ichthyosaursOmphalosaurus sp., and Shastasaurus sp. or proterosaur Tanystrophaeus conspicuus [7]. New skeletal reconstructions are based on the osteological analysis of three dimensionally preserved bones and skeletal remains. The large number of marine endemic placodont macroalgae feeders (P. gigas) in the Bayreuth sites coincides with the presence of invertebrate palaeocommunities that are characteristic of macroalgae meadow paleoenvironments. Most of the reptile species and genera from the Bayreuth localities also occur in beds of similar ages from the Monte San Giorgio (Switzerland/Italy) or Perledo (Italy) lagoonal areas. Ichthyosaurs and pistosaurs were adapted for open marine conditions, and may have migrated from the Panthalassa Oceans into the shallow marine Germanic Basin to reproduce, whereas placodonts and many other sauropterygians seem to have lived permanently in those shallow marine habitats, with large squamates and thecodont or smaller archosaurs in coastal areas.

During the last Ice Age of central Europe, cave bears hibernated deep inside their caves and Ice Age spotted hyenas used the cave entrance areas as dens. Steppe lions were also sporadic cave dwellers in order to hunt the herbivorous cave bears or steal prey from hyena dens, or for antagonistic and territorial reasons. Steppe lion skeletons recently discovered in the midst of cave bear skeletons, deep inside the Ursilor Cave (Romania), and steppe lion remains from other European caves, in particular a large population from the Zoolithen Cave in Germany, provide evidence of active predation by lions on cave bears. However, it was not mainly lions that fed on the cave bears: the large quantities of damaged cave bear bones, including incomplete long bones that repeatedly show similar types of damage, crushed long bones, and damaged skulls, reveal that hyenas were the primary scavengers on cave bear carcasses and were largely responsible for the destruction of their carcasses and bones. Predation and scavenging on cave bears (mainly by these two Ice Age predators but also by wolves and leopards) particularly in mountainous boreal forest environments that were devoid of the mammoth steppe fauna, explains the large quantities of fragmented cave bear bones that have been found all over Eurasia. The stress caused by these carnivores, deprived of their specialized ecological niche, may have led the herbivorous cave bears to hibernate as deeply as possible within their hibernation caves, in an attempt to protect themselves against predation. The felid predators were sometimes killed inside the caves, probably in conflicts with adult cave bears or even with hyenas, and remained as complete or only partly disarticulated carcasses.

The 22 meter thick marine carbonate Upper Oligocene series of Astrup (NW Germany) is correlated with the Chattian type section of Doberg. It indicates a more constrained palaeogeographical and biostratigraphical position ranging from the biozones of Chlamys (C.) decussata (upper Chattian A) to Chlamys (C.) semistriatus (lower Chattian C). The macrofauna can be subdivided into three main benthic communities: A. the „coarse gravel spondylid beach fauna“ of the shore zone with „pebble beach facies“ dominated by sessile brachiopods, large balanids, spondylids, oysters or small regular echinoids. Borings are common in pebbles; B. the „glauconite fine gravel brachiopod-bryozoan littoral fauna“ of the shallow subtidal zone where a terebratulid/lithothamnid dominated fauna/flora is present. The rhodophyceans were most possibly anker stones and substrates for cirripeds and serpulids; C. the „glauconite carbonate sand phytal fauna“ of the shallow subtidal zone with a rich benthic mollusc dominated fauna. Indirect evidence for seagrass and macroalgae occurs on the attachment negatives of balanids and oysters, and also on Cibicides foraminifera or bryozoans like Cellepora. The facies types along the Wiehengebirge Island and Teutoburger Wald Peninsula coasts of the southern Pre-North Sea Basin differ with respect to their benthic communities to that of the siliciclastic Leipziger and the Rhenish Bay facies.

One of the Europe's most complete Crocuta crocuta spelaea (Goldfuss 1823) skeletons from the Konĕprusy Caves, Bohemian Karst (Czech Republic) is an older pathological (hindlimb arthritis) and possibly smaller female individual. Few other cannibalistically scavenged hyena remains of older animals demonstrate a ‘small population’ that used the Main Dome area as a den site, which was marked well by faecal pellets. A large amount of megafauna prey was accumulated, and bones were further damaged or cracked. These hyenas hunted Przewalski horses dominantly, as has been documented all over the Bohemian Karst at several Late Pleistocene hyena den caves. The mammoth was mostly absent, instead the woolly rhinoceros was one of the meat colossi with indestructible bones also found in the Main Dome. These are the best indicators for hyena den sites identifications in general. Those massive bones found are left in repeating damaged ways and mainly as shafts and exposed best with bite marks in the bone spongiosa. Rhinoc...

The Upper Rhine Valley (SW Germany, Europe) has delivered 20,000 Late Pleistocene megafaunal remains found at gravel pit sites in the past decades and only 0.1% carnivore remains including a nearly complete lion skeleton of a small male Panthera leo spelaea (Goldfuss, 1810) and also skeleton parts of two more grown-up to even older males from different localities, such as the additional isolated bones. Less than 1% of the bone material belongs to adolescent animals; cub material is completely absent. The material from the Upper Rhine Valley was compared with many caves and few open-air sites of central Europe, indicating strong sexual dimorphism in the last lions of Europe, but problematic remains in the glacial/interglacial size variation. Some lions are in the open air, and all cave localities are overlapping with hyena and cave bear den sites; in some cases, they seem to have been in the open-air hyena den sites in the Upper Rhine Valley. Proofs for hyena or other large carnivore scavenging activities ...

The oldest record of Cyamodus is a skull of Cyamodus tarnowitzensis (Gurich 1884, Zt dt Geol Ges. 36:125–144) from Pelsonian shallow marine deposits. During the middle Illyrian, placodonts disappeared from the Germanic Basin. With renewed upper Illyrian transgression, Cyamodus rostratus (Munster 1839, Uber einige ausgezeichnete fossile Fischzahne aus dem Muschelkalk bei Bayreuth. Birner, Bayreuth, p. 14) appeared which was found in terebratulid shell-rich shallow marine deposits. Abundant remains of Cyamodus muensteri (Agassiz 1839), traditionally referred to as Cyamodus hildegardis and here synonymised with C. muensteri, have been reported from the upper Illyrian/middle Fassanian. Skeletal remains of this species are from the Grenzbitumenzone of the Monte San Giorgio lagoons. The youngest species, Cyamodus kuhnschnyderi (Nosotti and Pinna 1993, Compt Rend Acad Sci Paris. 317:847–850), has been found in the upper Fassanian/lower Longobardian of the southern Germanic Basin or Burgundian Gate when marine fa...

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 ...

The Middle to Upper Cenomanian (Upper Cretaceous) carbonate series of the Starigrad Limestone Formation outcropping on Hvar (studied sections equal 217 m) is characterized as six recurring facies associations that dominantly represent shallowing upwards parasequences comprising shallow marine/lagoonal through to supratidal flat strata. Facies Association 1 represents a shallow carbonate ramp–coastal environment patch reefs composed of rudists, other various bivalves and characteristic giant clams of unclear systematic position. Those giant clam/rudist patch reefs built benthic islands with other small oysters and spondylids in which a large number of small reef-dwelling Echinids lived. Facies Association 2 comprises platy limestones that are interpreted to represent low-energy lagoonal sedimentation. This lagoon paleoenvironment was at times exposed to hypersaline and anoxic bottom conditions, which resulted in the preservation of many marine vertebrate skeletons (fish and lizards) reported over the past two centuries. Only two different macroalgae, the calcareous Thaumatoporella, and a large currently unnamed probably non-calcareous form, are commonly observed within the platy limestones. Facies Associations 3 through 6 represent a complex range of hypersaline-associated strata that is dominated locally by anhydrite (Facies Association 3) and biolaminites of distal intertidal (Facies Association 4), middle intertidal (Facies Association 5), and upper intertidal/supratidal (Facies Association 6) affinity. Previously unrecognized are the intertidal Upper Cenomanian stromatolites and biolaminates at the top of the shallowing upward sequences. On those extended biolaminate-dominated sandflats dinosaurs migrated at minimum in the Mediterranean leaving extensive trackway systems at several places in Croatia.

Bone remains and a trackway of Pantheraichnus bottropensis nov. ichg. ichnsp. of the Late Pleistocene lion Panthera leo spelaea ( Goldfuss, 1810 ) have been recovered from Bottrop and other open air sites in northern Germany. Some of these bones are from open air hyena den sites. A relative high proportion of lion bones (20%) exhibit bite, chew or nibble marks, or bone crushing and nibbling caused by a large carnivore. Repeated patterns of similar bone damage have been compared to bone remains found at hyena dens in gypsum karst areas and cave sites in northern Germany. Ice Age spotted hyenas have been the main antagonists and the main scavengers on lion carcasses. The remains appear to have been imported often by hyenas into their communal dens, supporting the theory of strong hyena-lion antagonism, similar to the well documented antagonism between modern African lions and spotted hyenas. Most of the lion bones from the open air hyena den at Bottrop are probably a result of such antagonism, as are the rare remains of these carnivores found within large hyena prey bone accumulations along the Pleistocene rivers. The Emscher River terrace also has the largest quantity of hyena remains from open air river terrace sites in northern Germany. Their cub remains, and incomplete chewed prey bones from mammoths and woolly rhinoceroses, typical of hyena activity, underline the character of these sites as cub-raising and communal dens, where their prey was accumulated along the riverbanks in a similar manner to modern African hyenas.

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 100 m 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.

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 ...

A solitary articulated skeleton of a middle-aged and diseased Panthera leo spelaea lioness from the Eemian interglacial has been found amongst numerous articulated skeletons of Palaeoloxodon antiqu...

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, Acheulean) over the Middle Palaeolithic (Micoquian/Mousterian), and to the final Palaeolithic (Magdalenien) 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 dominantly 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.

A systematically excavated track site in a 243.5 Myr 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 40 m, 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, Biological Journal of the Linnean Society, 2011, 103, 76–105.

The remains of a large population of Late Pleistocene Ice Age spotted hyenas (Crocuta crocuta spelaea Goldfuss 1823) are described from the Rosenbeck 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 Rosenbeck 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, Biological Journal of the Linnean Society, 2011, 103, 191–220.

This paper presents a preliminary investigation of late Pleistocene cave bear traces from Ursilor Cave in the western Carpathians, Romania. The bears left thousands of traces on the walls, plateaus, and slopes of the cave interior. Some areas in the cave have been heavily trampled, leaving more than 140 hibernation beds as well as fur impressions. The footprints of cave bears are assigned to Ursichnus europaeus nov. ichnogen. and nov. ichnosp. and the cave bear-beds to Ursalveolus carpathicus nov. ichnogen. and nov. ichnosp. as behavioral traces. Tens of thousands of scratch marks on the slopes and top of a clay plateau are the result of bears moving from the hibernation area down to a stream and back. These traces reveal that the cave bears had short claws, similar to those of modern, primarily herbivorous black bears. Deep within the cave, three weathered, articulated cave bear skeletons still lie in their hibernation beds on the clay plateaus or a natural cave corner. One of these bears was a one-year-...

A femur fragment with an Early Lutetian (early Middle Eocene) age is the world’s oldest fossil record from a seal, and, is described as Praephoca bendullensis nov. gen. nov. spec. This find pushes back the earliest evolution of seals into the Paleocene epoch. The femur has plesiomorphic terrestrial mammal characteristics but has a morphology that is already closer to that of Miocene and present day seals. The Eocene seal femur was found at Furstenau-Dalum in north-west Germany, in a conglomerate rich in shark teeth that was deposited in a coastal delta environment to the north-west of the central European Rhenish Massif mainland, in the southern pre-North Sea Basin. This discovery has led to a revision of the theory that phocids originated along the coastline of the North American continent. Instead they can now be interpreted to have originated in the tropical Eocene climate of central Europe. Although the fossil records of pinnipeds in Europe during the Eocene, Oligocene and Miocene are extremely sparse, they appear to have inhabited the pre- North Sea basin, within the influence of temperate and arctic upwellings. The distribution of abundant teeth from white and megatooth sharks of two different lineages appears to correlate with that of the seals, which the sharks most probably hunted; providing supporting evidence that the phocids were already adapted as shallow marine coastal inhabitants by this time.

The descriptions of fossil Triassic marine pla- codonts as durophagous reptiles are revised through comparisons with the sirenia and basal proboscidean mammal and palaeoenvironment analyses. The jaws of placodonts are conver- gent with those of Halitherium/Dugong or Mo- eritherium in their general function. Whereas Halitherium possessed a horny oral pad and counterpart and a special rasp-like tongue to grind seagrass, as does the modern Dugong, placodonts had large teeth that covered their jaws to form a similar grinding pad. The sirenia also lost their anterior teeth during many Millions of years and built a horny pad instead and specialized tongue to fed mainly on seagrass, whereas placodonts had only macroalgae availa- ble. Indirect evidence for Triassic macroalgae is provided by benthic palaeocommunities from different layers and extended European regions in the Germanic Basin. Studies of tooth wear stages for Placodus indicate that anterior teeth may have been used in a similar manner to the procumbent front teeth of modern Dugong. Paraplacodus and Placodus seem to have used these teeth as spatulas to dig out seaplants. Cyamodus and other placodonts such as Placochelys had smaller or reduced anterior teeth. The scarcity of highly worn palatine or maxillary and lower jaw dentary Placodus or Cyamodus teeth (less then 0.5%) suggests that they had a relatively soft diet. The seaplants would only have been squeezed in a similar feeding strategy to that of modern Dugong feeding on seagrass without jaw rotation and grinding. The phylogenetic trend in tooth reduction within the placodonts Paraplacodus, Placodus, especially in Cyamodus but also Placochelys, and Henodus within 11 My appears to have been a result of this plant-feeding adaptation and may even explain the origin or at least close relationship of the earliest Upper Triassic turtles as toothless algae and jellyfish feeders, in terms of the long-term convergent development with the sirens.

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".

Wolverine bone material is described from the famous Upper Pleistocene cave Srbsko Chlum-Komin in the Bohemian Karst, Czech Republic, along with an overview of recently known Czech sites. The Gulo gulo Linnematerial was found in one of the largest Ice Age spotted-hyena dens in Europe. As a result of non-systematic excavations, the taphonomy is partly unclear. Lower-jaw remains indicate a minimum of three wolverines. Two of the mandibles are cracked, which is most likely the result of carnivore scavenging. The absence of juvenile G. gulo suggests possible importation of the wolverines by hyenas Crocuta crocuta spelaea Goldfuss.

Late Pleistocene spotted hyenas hunted Przewalski horses in spring to early summer, as documented by foetal horse skeleton remains of Equus ferus cf. przewalskii Poljakoff 1881, found between 3569 megamammal bones in the hyena den site Srbsko Chlum-Komin of the Bohemian Karst (Czech Republic). The main prey of hyenas Crocuta crocuta spelaea (Goldfuss 1823) was this small horse, well distributed in Central Europe. 51% (NISP = 1.800) of the megafauna in the Srbsko Chlum-Komin Cave are horse bones, consisting of mainly complete and in a few cases cracked and chewed dominantly leg and cranial remains. Carcass body parts must have been imported into the hyena prey storage and cub-raising den cave. Similar high amounts of horse remains and taphonomic situations are reported from other Late Pleistocene hyena den caves in Central Europe, such as Rochelot Cave (France). Den sites indicate an active selected hunt of mainly adult horses. Hyenas left partly articulated legs or complete distal bones and many isolated teeth at their dens, the latter resulting from consumption of the thin-walled horse skulls. In periods of megafauna species abundance, hyenas used mainly the meat leaving only a few chewed or cracked horse bones (Srbsko Cave or Rochelot Cave), whereas in poor feeding times or in cub-raising dens, bone consumption was higher resulting in many fragmented and chewed horse bones (Nad Kacakem Cave, Czech Republic). Whereas African spotted hyenas hunt zebras and even modern donkeys successfully, in comparison to modern spotted hyena dens in Amboseli (Kenya) and Syokimau (Botswana), it seems as if the Ice Age spotted hyena Crocuta crocuta spelaea of Europe hunted very similar equine prey, in this case the Przewalski-horse, especially in hilly and mountainous regions, where the largest prey (mammoth and woolly rhino) were rare or absent.

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupus spelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.

Upper Pleistocene remains of the Ice Age steppe lion Panthera leo spelaea (Goldfuss, 1810) have been found in the Perick Caves, Sauerland Karst, NW Germany. Bones from many hyenas and their imported prey dating from the Lower to Middle Weichselian have also been recovered from the Perick Cave hyena den. These are commonly cracked or exhibit deep chew marks. The absence of lion cub bones, in contrast to hyena and cave bear cub remains in the Perick Caves, and other caves of northern Germany, excludes the possibility that P. leo spelaea used the cave for raising cubs. Only in the Wilhelms Cave was a single skeleton of a cub found in a hyena den. Evidence of the chewing, nibbling and cracking of lion bones and crania must have resulted from the importation and destruction of lion carcasses (4% of the prey fauna). Similar evidence was preserved at other hyena den caves and open air sites in Germany. The bone material from the Perick and other Central European caves points to antagonistic hyena and lion conflicts, similar to clashes of their modern African relatives.

More than seventy-five vertebrate track-sites have been found in Central Europe in 243–246.5 m.y. old Triassic coastal intertidal to sabkha carbonates. In the western part of the very flat Triassic intracontinental Germanic Basin, the carbonate strata contain at least 22 laterally extensive track horizons (called megatracksites). In contrast, in the eastern part of the basin only six megatracksites extended to near the centre of the Basin during marine low stands. Marine ingression and the development of extensive coastal marine environments began during the Aegean (Anisian) stage. This incursion began in the region of the eastern Carpathian and Silesian gates and spread westward due to the development of a tectonically controlled intracratonic basin. The tectonic origin of this basin made it susceptible to tsunamis and submarine earthquakes, which constituted very dangerous hazards for coastal terrestrial and even marine reptiles. The shallow sea that spread across the Germanic Basin produced extensive tidal flats that at times formed extensive inter-peninsular bridges between the Rhenish and Bohemian Massifs. The presence of these inter-peninsular bridges explains the observed distribution and movement of reptiles along coastal Europe and the northern Tethys Seaway during the Middle Triassic epoch. Two small reptiles, probably Macrocnemus and Hescherleria , left millions of tracks and trackways known as Rhynchosauroides and Procolophonichnium in the Middle Triassic coastal intertidal zone. The great abundance of their tracks indicates that their trackmakers Macrocnemus and Hescherleria were permanent inhabitants of this environment. In sharp contrast, tracks of other large terrestrial reptiles are quite rare along the coastal margins of the Germanic Basin, for example the recently discovered archaeosaur tracks and trackways referable to Isochirotherium , which most probably were made by the carnivore Ticinosuchus . Smaller medium-sized predatory thecodont reptiles, possibly Euparkeria , probably made the Brachychirotherium trackways that have been found across much of Central Europe. Large lepidosaurs such as Tanystrophaeus probably hunted in the tidal ponds and channels, where they locally produced Synaptichnium tracks. Recently discovered tracks made by a basal prosauropod are the world's oldest record of this group of dinosaurs, occurring in beds that have an age of about 243.5 Ma. (Pelsonian substage). This shows that very large prosauropods existed much earlier than was previously believed. These prosauropod tracks, along with tracks of small bipedal dinosaurs found in the Alps and Eastern France, show that by the middle part of the Middle Triassic the radiation and diversification of dinosaurs was already in progress. In the Germanic Basin, aquatic-adapted paraxial swimming sauropterygians are not known to have left tracks, except for occasional subaquatic swimming scratch-mark “trackways” within the coastal tidal flat zone. Marine-adapted aquatic reptiles migrated into the Germanic Basin with increasing frequency in the upper part of the Middle Triassic, when the bathymetry of the Germanic Basin was at its deepest following a strong regression that occurred due to basin uplift in the middle part of the Middle Triassic. These large marine reptiles included Pistosaurus , the ichthyosaurs Cymbospondylus or Mixosaurus , and many placodonts such as Cyamodus , Placodus and Paraplacodus , which fed on macroalgae and seem to have been the Triassic sea cows of their day. The distribution of these reptiles was mainly controlled by tectonics, but eustatic changes in sea level also were important and produced widespread environmental changes across the tidal flats up until their disappearance in the Germanic Basin in the late Middle Triassic. The initial break-up of Pangaea already had started in Middle Triassic time, and this event had begun to drastically change environments all over Central Europe. It is very interesting that dinosaurs began to diversify at exactly this time, and it seems likely that this was a direct reaction to these environmental changes. It can be inferred that the earliest dinosaurs must have started to evolve in the late Early Triassic, because in Europe it can be demonstrated that at least two main dinosaur groups already were present and clearly differentiated by the middle part of the Middle Triassic, and all three of the major groups of dinosaurs (theropods, sauropods and ornithischians) had diversified and spread globally throughout terrestrial habitats by the end of the Triassic Period. Six new palaeogeographic maps, representing time intervals from the Aegean to the Illyrian (Anisian) stages, show these important environmental changes in detail and explain the direction and timing of terrestrial reptile exchanges between the Central Massif, Rhenish Massif, and Bohemian Massif, and also the direction and timing of marine reptile exchanges between the Alps of Central Pangaea and the ancient northern Tethys Ocean and Germanic Basin Sea.