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Recycling of badger/fox burrows in late pleistocene loess by hyenas at the den site Bad Wildungen-Biedensteg (NW, Germany): woolly rhinoceros killers and scavengers in a mammoth steppe environment of Europe.

1. Introduction

Late Pleistocene European bone assemblages have been produced mainly by late Ice Age spotted hyena Crocuta crocuta spelaea [1] and were first recognized by Buckland [2] in the "Kuhloch Cave" (Konig-Ludwigs Cave, Bavaria, Germany) and the Kirkdale Cave (Kent, England). More recent studies provide information on the hyena prey bone assemblages (e.g., [3-10]) as well as on the new subdivided fossil hyena den types (e.g., [11]). These identifications of three classified Ice Age den forms are particularly important also to distinguish bone accumulations made by hyenas from those accumulated by Middle Palaeolithic humans (e.g., [9,12-15]).

Few contemporary used hyena and Neanderthal sites have been described from hyena dens in mammoth steppe lowlands and adjacent cave-rich region environments of north-central Europe, in England and Germany [9, 16]. The degree of prey bone damage and presence/absence of "nibbling sticks" and faecal pellets or hyena population structure and their individual amount allow the reconstruction, much better, of the ethology of the last hyenas of Europe. The discussions for nonarchaeological sites no longer focus only on the human/carnivore origin discussion. Although hyena cave-den sites predominate in the European fossil record (e.g., Germany in [17]), open air sites may have been much more common throughout the mammoth steppe lowlands of Europe, but have been overlooked or not identified as such (cf. Westeregeln or Bottrop sites in [10,18]).

Open air hyena den sites in loess deposits without human impact are not analyzed in Germany, as yet, whereas other bone accumulation sites on river terraces have been analyzed along the Emscher River near Bottrop in the Westphalian mammoth steppe lowland [10]. Recently many open air hyena den sites (loess, gypsum karst, river terraces: Saalfeld, Bottrop, Westeregeln, Sewecken-Berge, Thiede, and others) from Germany have been described [17, 19-21], whose density overlaps with the Middle Palaeolithic Neanderthal occupation and open air and cave sites in Germany, even in the famous Neanderthal valley [22, 23]. Additionally, the review of lion localities in northern Germany [24] demonstrates not only quite hard competition conditions about megafauna prey between those two top predators killing and consuming each other, but also competition with human Neanderthals during the Late Pleistocene. In Germany, additionally, mostly hyena den cave sites have been described and newly identified, also partly overlapping with human camp sites, for example, Balve Cave [17, 22, 23, 25-27]. The herein reviewed hyena den site Bad Wildungen-Biedensteg is not far from a Middle Palaeolithic site Buhlen (Micoquien to Late Mousterien: [28]), but has no evidence of human impact.

History of the Bad Wildungen Hyena Den Site. First Ice Age fauna remains in the clay pit site "Ziegeleigrube Biedensteg" in Bad Wildungen-Biedensteg of northern Hesse (Central Germany, Figure 1, GPS coordinates: long. 9[degrees]8,24.32"E, lat. 51[degrees]7716.44"N) were discovered in 1932 by the hobby paleontologist/archaeologist Pusch, who excavated and rescued many macromammal bones. In 1952 Jacobshagen and Lorenz found a micromammal-rich "pellet horizon" and two hyena skulls [29]. Jacobshagen described in 1963, briefly, this fauna, but wrote mainly about the micromammals. Huckriede and Jacobshagen [30] published the first section, which was studied with an addition of new sedimentological results by Semmel [31] and Kulick 32]. The last micropalaeontological research was performed by Storch [33] on pellet material. First thoughts about hyena gnawing and bone deposits were mentioned by Jacobshagen [34] with new research being published about the hyenas, woolly rhinoceros, and cave bears [35]. Here, the complete megafauna and hyena den site analyses are presented in more broad comparisons to many other new analyzed Late Pleistocene hyena dens studied these past years in Germany and Czech Republic (Figure 1).

2. Material and Methods

The main collection (including coll. Pusch, coll. Lorenz) is owned by the Rudolf-Lorenz-Stiftung (coll. no. Bi-52/1237) and was partly presented in the "Stadtmuseum of Bad Wildungen." Additionally, a few macromammal bones from the collection in the "University of Marburg" were integrated in this study, which was also mentioned in the article of Jacobshagen [34]. This collection was partly rediscovered by Dr. Fichter, who kindly helped by donating the important micromammal collection to the "Kurmuseum Bad Wildungen." Only Kulick [32] made a small systematic excavation at the site, which produced mainly micromammals from pellets.

Comparative bone material was used in many different collections. The most important is the woolly rhinoceros skeleton from Petershagen (NW-Germany) in the Museum Natur und Mensch Bielefeld (MNMB). Another mounted skeleton cast in the Museum fur Ur- und Ortsgeschichte Eiszeithalle Quadrat Bottrop (EMOB) was used for the skeleton redrawing and comparison of the bone positions in the skeleton of the Bad Wildungen-Biedensteg material. Skeletons of the extinct Przewalski horse (Equus caballus przewalskii) were studied in the Julius-Kuhn Museum Halle/ Saale (JKMH; see also [36]), reindeer (R. tarandus) and arctic fox (V lagopus) skeletons in the collection of the University of Alberta Department of Biological Sciences (UADBS); mammoth (M. primigenius) remains and cave bear (U. spelaeus) and red fox (V. vulpes) bones were compared to skeletal material in the Geologisch-Palaontologische Museum der Westfalischen Wilhelms-Universitat Munster (GPIM). Finally, recent badger (M. meles) or common hare (L. europaeus) and the Pleistocene hyena materials from the Srbsko-Chlum were used in the collection of the National Museum Prague (NMP) and from the Perick Caves of the Staatliche Naturhistorische Sammlungen Dresden (SNSD). The open air gypsum karst site Westeregeln material was studied in the Martin-Luther-University Halle/Saale (MLU.IFG) and the Natural History Museum of the Humboldt-University Berlin (MB).

3. Sedimentary Geology,

Paleoenvironment, and Dating

The geological situation at the hyena site "Lehmgrube Biedensteg" was published by Huckriede and Jacobshagen [30], Semmel [31], and Kulick [32]. The overview of the redrawn sketch of the outcrop section, with a combination of all published results and new interpretations about the hyena deposits, is presented in Figure 2.

The Wilde River gravels at the base of the section are of the Eemian Interglacial period. They consist of Red Bunter sandstone and claystone, lydite, quartz, or diabase pebbles. These deposits are overlain by a palaeosoil resulting from solifluction. In this "Eemian Soil" the river pebbles are resedimented with reddish-brown loess. The "Lower Loess" is from the early to middle Lower Weichselian (MIS 5c-d), and after Semmel [31], a product of the first part of the glaciation (early Late Pleistocene, Figure 2), where, in this mountainous region, loess was deposited in a mammoth steppe environment. Some snails were found in the Lower Loess by Jacobshagen [34], the mentioned loess soil snail Pupilla muscorum (Mualler) fitting to the cold period climatic and environmental mammoth steppe interpretation.

In the middle and at the end of the Late Pleistocene a climatic stagnation resulted in a palaeosoil along the Wilde River gravels which were, at that time, on the shore of a small lake. This lake was caused by subsurface salt dissolution and positioned in a large-scaled sinkhole structure. The lake was filled up by the Wilde River, indicated by the presence of many aquatic vertebrate species, such as frogs (Rana agiloides Brunner), but mainly by salmonid fish (cf. [34]) that lived in fluent water.

The muddy area at the Wilde River or lake shore was used by the Ice Age spotted hyenas as prey deposit sites [35]. Bones from animals of the mammoth steppe macrofauna were deposited here, whereas "bone nests" were mentioned in the publication of Jacobshagen [34]. The sedimentary depression structures in the bone-rich loess horizon described by Kulick [32] as "cryoturbation and channels" also could be partially of bioturbation origin and were possibly caused by the hyenas who deposited animal prey remains in the soft soil, only in summer times, when the permafrost soil was soft in the upper parts.

The bioturbation interpretation would fit into the "hyena commuting/prey storage site," but can no longer be studied because of the nonopen loess pit Biedensteg. In this section (Figure 2) such depressions are figured as hyena prey depots. Possibly, a later cryoturbation, a result of permafrost soils fitting into the environment and climatic situation of that time, was responsible for secondary overprint of the primary sediment structures. Bioturbation by mammoths on lake shores, which left depressions of their footprints, must be taken into account, as is discussed for other sites (cf. [37]).

The "pellet horizon" is figured differently in the publications (of Jacobshagen et al., 1963, [32]). The section of Kulick [32] indicates that the pellets and the macromammal bones are mixed in a single horizon. Proof for that might be caliche concretions around hyena coprolites in which micromammal bones and teeth are also cemented in. The "hyena prey depot site" and the "pellet horizon" are from the same period and are dated relatively (no absolute data) into the late Middle Late Pleistocene or Weichselian (65.000-90.000 BP, MIS 5cd, Figure 2).

The bone-rich horizon is overlain by another palaeosoil, the "Lohner Soil," which can be found in the region at different sections [31, 32]. After their interpretations a solifluction of Loess and Wilde river gravel material took place in the middle Late Pleistocene warm period (Figure 2). VI vulpes and M. meles were the dominating faunal elements, besides L. europaeus. This fauna fits to Meles/Vulpes den burrow sites in loess soils, in front of which they often left some prey bones.

Finally the upper loess was deposited within the LGM, and after, the upper part was decalcified during the Holocene period. The "Eltviller Tuff" is a one to two centimeter thin layer in the upper loess and the only absolute dated horizon with an age of around 16.000 BP ([31], Figure 1(c)).

4. Small Carnivore Fox Den and Mustelid Bone Assemblage

Meles meles (Linne 1758) (Figure 4(13)-(32)) (Table 5) is known by one skull of an adult male (Figure 4(13)) and a second brain case of a juvenile. Several postcranial bones consist of the forelimb (Figure 4(14)-(20)) and hind limb bones (Figure 4(21)-(29)), although vertebrae are missing (cf. Table 1).

Vulpes vulpes (Linne 1758) (Figure 4(1)-(9)) remains consist of 13 common fox bones (Table 3) including a skull. This skull is incomplete, as most of the anterior part with its dentition is missing. The last three teeth are in the left maxillary (Figure 4(1)). From a right forelimb the scapula, humerus, and radius were found, which seem to belong to one individual (Figure 4(2)-(4)). From a hind limb, not only the left femur shaft and incomplete tibia but also a right calcaneus and a metatarsus III are represented (Figure 4(5)(8)). A fragment of a metapodial is missing its proximal joint. Finally a lumbar vertebra and one rib are preserved. The pelvis is missing its left part (Figure 4(9)). A second pelvis fragment is again incomplete. Material from two individuals is present, indicated by the pelvis remains. Possibly most of the bones belong to only one individual. All postcranial bones show a complete fuse of the symphyses and are from either a single animal or several adult animals.

Vulpes lagopus (Linne 1758) (Figure 4(10)-(12)) (Table 4) was found with a nearly complete skull, without the jugal arches, but with the right mandible (Figure 4(10)-(11)). The skull sutures are not fully fused and teeth are barely used; therefore it was a young adult individual, as only a single individual can be estimated from the bone material. The postcranial material is present with a femur shaft and pelvic fragment (Figure 4(12)).

Mustela putorius Linnaeus 1758 (Figure 4(33)) (Table 6) is present with a single half skull (Figure 4(33)) of which the anterior part with most of the dentition is preserved.

Lepus europaeus/timidus Linne 1758 (Figure 14(1)-(9)) (Table 14) is represented by 28 bones which are cranial fragments, two are mandibles and the rest are postcranial bones (Table 13). There is an articulated pedal skeleton (Figure 14(9)) and an articulated pelvis with lumbar vertebral column (Figure 14(5)). The figured material (Figure 14) seems to be from one individual, which is indicated by the bone preservation and articulations. Another argument is the individual adult's age and the fresh fractures of the humerus, radius, the right femur and left tibia, or some processes of the vertebrae, which were caused during the excavations. Bones from other individuals of young and adult age are also preserved and have been completely disarticulated. 25% of the remains are from young animals; 75% are from adult hares. Three animals can be estimated by the tibia as minimum individual number.

5. The Hyena Population and Coprolite Remains

The Ice Age spotted hyena Crocuta crocuta spelaea [1] (Figure 2) skeletal remains consist of four skulls, three mandibles, one radius, and a femur (Table 1). Additionally, there are 16 coprolites which were rescued.

From the first skull (Figure 2(1)) deformations do not allow exact metric data. The second skull (Figure 2(2)) is 290 mm in total length and measures 265 mm between the incisive and condyle. The largest height is behind the frontal processes (114 mm). The distances between the canines and P4 are about 68 mm. The width of the frontals (zygomatic processes) measures 90 mm. Finally the outer distance between the canines is 58 mm. The largest diameter of the canines in the middle of the tooth is 18 mm. The brain case symphyse of the third animal (Figure 2(3)) is slightly fused and articulated. The parietal, frontal, palatine, and temporal are incomplete. The maximum width measured, between the temporal, 73 mm, whereas it is preserved in 76 mm in length.

One left mandible (Figure 2(4)) is of an adult animal and might belong to one of both individual adult skulls, which show a similar tooth use stage. The jaw was cracked by hyenas between the [P.sub.2] and [P.sub.3]; the [P.sub.3-4] and M3 are present. The ramus was damaged during excavations.

A few postcranial bones are represented with one axis of an adult animal exposing bite damage marks (Figure 2(6)). A left radius and a left femur (Figure 2(5) and (7)) are from one very young cub, both being incomplete as a result of scavenging activities by large carnivores.

Coprolite Material. The hyena coprolites are generally white inside and the pores are filled with iron and manganese minerals. The coprolites show a moderate variability and even bone contents (Figure 2(8)-(17)). The largest one (Figure 2(8)) is a double pellet being connected by caliche incrustations. It seems to represent a fossilized, originally softer and humid, faecal pellet. The other pellets have repeating shapes and have attached 3-5 smaller pellets (Figure 2(9)-(12)), representing possibly more dry dung. Single pellets have often defined shapes. The most represented one is the "drop shaped pellet" (Figure 2(13)-(15)). They can point to both sides or can end round to flat on one side as a result of attachment to another pellet. Other pellets are "unshaped" and irregular. These were often found in the non-spindlelike pellet aggregations (Figure 2(10)). In the material from Biedensteg each coprolite contains several bone fragments, which are often visible on the surfaces (Figure 2(11)-(12)). These are small pieces, well rounded by stomach acid, and are mainly from the bone compacta, but also are isolated pieces of bone spongiosa. This spongiosa is very thin walled and should have been completely dissolute. These spongiosa pieces are most comparable to the bone spongiosa of the woolly rhinoceros, but might also refer to other megamammals.

6. Hyena Megafauna Prey Remains

Ursus spelaeus Rosenmuller 1794 subsp. (Figure 3) is represented by four cave bear bones and fragments. The left scapula (Tables 2 and 3(1)), which lacks all distal parts seems to be destroyed by hyenas. Large carnivore gnawing and bite marks are visible at the glenoid. A right humerus shaft (Figure 3(2)) is missing the joints as a result of heavy carnivore chewing. At the shaft ends and in the lower middle, bite marks are present. The diameter of the bone shaft is small, being only 49 mm. From one left incomplete ulna (Figure 3(3)) the distal joints were chewed and also some bite marks are visible. The 50 mm maximum width ulna has, again, small proportions. Finally, a fragment of a femur shaft (Figure 3(4)) with heavy chewing damage indicate the cracking and further use of the bone fragment as a typical hyena "nibbling stick" (for teething purposes of hyena cubs).

Mammuthus primigenius (Blumenbach 1799) (Figure 12 (1)-(3)) is represented by three remains consisting of a tooth lamella fragment from a juvenile animal, a thoracic vertebra neural arch and centrum fragment, and a long bone fragment used as a nibbling stick (Table?). The material is from adolescent elephants.

Coelodonta antiquitatis (Blumenbach 1799) (Figures 5-11) is the most abundant, listed in Table 8. The cranial elements consist of a middle part of a skull from a young calf (Figure 9). The connection in-between the maxillas were restored in former times. Originally, the maxillary part between the teeth was damaged by hyenas. All three [dm.sup.1-3] milk teeth on both sides are present (Figure 6(1a)-(1d)). Both [m.sup.1]'s are breaking through, whereas the [m.sup.2]'s were still in the maxillary. These are not present, but the alveolar grooves are preserved. This skull was badly damaged by the hyenas, especially at the anterior part and the brain case. The latter shows a very interesting large carnivore brain case opening. There are some bite marks, but thin parallel long scratch marks on the right maxillary in the high of the [dm.sup.2-3] could have resulted from other smaller carnivores or hyena cubs. Both mandibles of the lower jaw (Figure 6(1e)-(1h)) fit to the skull by the identical milk dentition of the [dm.sup.1-3] and the tooth rising of the [m.sub.1]. Both jaws were cracked in the symphyses area and have old fractures. Additionally, they are lacking the rami and have large carnivore chewing and gnawing marks (Figure 6(1e)-(1h)). The left jaw possesses the [dm.sub.1-3] and the [m.sub.1]. The right mandible was damaged by the excavations and because of this is lacking the anterior part, including the [dm.sub.1-2]. Other cranial material was described and partly refigured by Jacobshagen [34]. He refigured some lower jaw teeth of one individual (right [P.sub.3-4], [M.sub.1], and left [M.sub.2-3]). The little use of the M3 indicates an origin of an early adult animal. It is suggested here that these belonged most probably to the skeleton of an early adult female individual (Figure 5(b)). Scapulae are preserved with one nearly complete left shoulder blade (Figure 7(1)). Some parts from the left side and joint area, destroyed by the excavations, were restored. Bite marks were found only distally. Here, hyenas left typical chewing marks in the very soft scapula. The margin is therefore typically irregular, resulting from cracked bone material. The scapula seemed to belong to the female skeleton. A second fragment of a scapula is in preservation and could be found in a lower horizon. One humerus is described by Jacobshagen [34], which can no longer be located. It was a right humerus that was chewed on the proximal joint. Ulnae are present with five bones (Figure 8(1)-(4)) from different old animals. The most juvenile, a neonate to young, animal's left ulna must have been articulated to one radius (Figure 6(2)). This result is from the comparison to an articulated right ulna/radius from a young adult to adult animal whose joints are chewed away in the same way (Figure 8(1)). The latter might belong to the young adult female rhinoceros (Figure 5(b)), of which also other bones were found partly articulated. At least seven radii (Figure 7(4)-(6), MNI = 7) were found, of which four are from young adult to adult animals and the last from the neonate to very young individual. The four pelvis remains are typical rests of hyena feeding activities (Figure 10(1)-(3)). The acetabular and surrounding two acetabular fragments are from different animals. The one figured (Figure 10(1)) has not only hyena, but also arctic fox, wolf or hyena cub, and even small rodent nibbling marks. The fourth pelvis remain is only a part of the ileum (Figure 6(3)) and seems to belong to the juvenile animal, because it is also chewed from the acetabular region. It is also heavily chewed at the soft distal part with irregular margin. Four femora are preserved, of which one is a fragment, a second is from a juvenile animal (Figure 6(4)), and a third and fourth are from an adult C. antiquitatis (Figure 10(4)-(5)). Another fragment is of an adolescent, with strong chewing marks (Figure 10(6)). As described by Jacobshagen [34], there was a right femur (Figure 10(4)) found in articulation with a tibia (Figure 11(2)). Only one nearly complete left patella (Figure 11(9)) was excavated and might belong also to the female skeleton's hind leg (Figure 5(b)). The tibia has very typical hyena caused damages and is in an early stage (stage 1) of destruction. Also this fits well with the partly articulated female skeleton carcass. Three tibiae are very massive and have a strong width in the shaft (Figure 11(3)-(5)). All tibiae compared indicate a sexual dimorphism with males being stronger and more massive in their bones. Mostly the proximal joint was chewed away first, although at the distal part in a middle stage (stage 2 of three) of bone feeding, two groves were left, which is documented at all three tibiae (Figure 11(3)-(5)). Two fibula remains are in the material, with one (Figure 11(7)) being proximally incomplete as a result of the excavations. That one was articulated to one tibia in the stage of hyena chewing and seems to belong to the female carcass (Figure 5(b)). The distal part shows long bite scratches. The second fibula was cracked away from a tibia and was left with the middle shaft with bite marks at both ends (Figure 11(6)). Only one astragalus and calcaneus are in the material (Figure 11(8)) also most probably belonging to the hind leg of the female skeleton (Figure 5(b)). They fit perfectly together, indicated additionally by overlapping bite scratch marks which are crossing both bones. After the descriptions by Jacobshagen [34] there were three complete metatarsals (2-4) that also fit for the female skeleton (Figure 5(b)), although it is unclear whether they are from the right or left side. All vertebrae show the typical hyena chewing by the lack of nearly all processes. They seem to be all from one nearly adult individual, indicated by a series of articulation and the similar degree of nonfusing of the caudal vertebra centrum disc. The cranial disc, in contrast, is already fused completely at all vertebrae. From the vertebral column, the first three cervical vertebrae were found connected (Figure 9(4)). Atlas (Figure 9(1)), axes (Figure 9(2)), and the third cervical vertebra (Figure 9(3)) have bite marks on the damaged processes. The next articulated vertebral column part is the vertebra from the sixth cervical to the first thoracic (Figure 9(7)). Articulated cervical vertebrae no. 6 (Figure 9(5)) and no. 7 (Figure 9(6)) and thoracic vertebra no. 1 (Figure 9(8)) are also lacking most of their processes, especially the dorsal ones. Two more articulated vertebrae are the second (Figure 9(9)) and third (Figure 9(11)) thoracic vertebrae which are heavily chewed (Figure 9(12)). The fourth thoracic vertebra (Figure 9(10)) was only a centrum that was found in nonarticulation with other vertebrae. The complete neural arch was eaten. Parts of the left side were cut by excavation activities. The longest articulated vertebral column part exists from the sixth to ninth thoracic vertebrae (Figure 9(17)). Typical for the hyena scavenging activities are the chewed dorsal spines. Finally, the articulated last thoracic and first lumbar vertebra were found connected (Figure 9(18)). Also, the first lumbar vertebra is lacking parts of the proc. transversus. The ribs generally have no hyena bite marks, but obviously they were removed from the carcass (Figure 9(20)-(28)). All costae have cracking fractures at both ends; all joints are lacking. Only one small rib fragment (Figure 9(28)) has distally small bite marks. Nibbling by a small carnivore, such as a young hyena, wolf, or arctic fox, has caused a pointed distal end. A small fragment was used for nibbling by young hyenas ("nibbling stick" no. 3, Figure 9(27)). The present rib fragments are from the anterior part around the forelimb, and a few are from the last thoracic vertebrae.

Bison/Bos (Figure 12(4)-(9)) remains consist of 13 bones (Table 9), two of which are teeth, the others being postcranial bones, which are all incomplete as a result of large carnivore activities. Most bones are limb bones, especially from the hind limbs. The teeth are two M1's, one from the upper and the other from the lower jaw. The strong tooth use indicates an individual of adult to older adult age. From the forelimb a metacarpal fragment (Figure 12(4)) was found. The metacarpal shows a typical hyena cracking preservation; the distal part has sharp edges. Most bones are from the hind limbs. Both femora were cracked in the middle of the shaft but also the distal joints were heavily eaten and nibbled (Figure 12(5)-(6)). One middle shaft of a cracked tibia and one proximally chewed calcaneus (Figure 12(8)) and two femur fragments seem to originate of the rigth hind limb of one animal. Finally, there is one thoracic vertebra centrum (Figure 12(9)) and one cervical vertebra (Figure 12(10)). The processes were chewed, and also some deep scratch bite marks can be found ventrally. All bones belonged to one, or possibly a few adult individuals.

Equus caballus przewalskii Poljakoff 1881 (Figure 13(4)-(15)) consists of 19 bones, of which two are mandible fragments, one cranial fragment and a single tooth, although mainly leg remains are represented (Table 10). The one metacarpus is 236 mm in length and distally 50 mm in width (Figure 13(8)) and falls within the small Przewalskii horse metapodial osteometry (cf. [9-11, 18, 37-44]). The same is for one complete metatarsus (Figure 13(15)) with its 257 mm length and 53 mm distal width. Also, there is the nearly complete lower jaw of a male horse (Figure 13(4)), as well as other small-sized bones from the smaller Przewalskii horse. There are bones from young horses (21%), with all others being from adult individuals (79%).

Megaloceros giganteus (Blumenbach 1799) (Figure 13(1)) was found with only seven bones, including one mandible fragment and three teeth, all from adult animals (Table 11). The material described and figured from Jacobshagen [34] is lost.

Cervus elaphus Linne 1758 (Figure 13(2)-(3)) is present with only two remains (Table 13). From the cranium, a right maxillary fragment with two [M.sup.1-2] shows the [M.sup.2] not in a developed state, although, the [M.sup.3] alveolar is opened and the tooth is in change. Another remain is a metatarsus (Figure 13(3)). All remains are from possibly a single calf, approximately 1.5 years old.

Rangifer tarandus Linne 1758 (Figure 13(16)-(25), Table 12) is more common, with 24 remains. The rest of the bone material, such as a right metatarsus, a phalanx 1 and phalanx 2 proximal joint disc, and a right radius distal joint fit in the nonfusing of the joints to one young animal. The dropped antlers are from males and are all from sheds, which must have been collected by hyenas. Similar damages are present on the distal ends where large triangular-oval bite impact marks and elongated scratches indicate large carnivore damage (Figure 13(15)-(17)).

7. Discussion

7.1. The Badger/Fox Types and Den Micromammals and Pellet Accumulators. At open air badger den sites, typically, most skulls and massive long bones were found, although such long-term used badger loess den systems are described [45]. In those, bone accumulations are dominated by skull remains, being figured, for example, for the Schneehalle Cave (South Germany, [46]). Commonly, badgers die in their dens [46-48], explaining their bone accumulations in burrows and caves. The amount of bones, mainly of senile and very young badgers of Bad Wildungen, fit into such a scheme. Bite marks and missing joints in a humerus and tibia might be the result of badger cannibalism [47] or even hyena activities. The skull and postcranial material can be referred to the Asian species Meles meles cf. leucurus (cf. [49, 50]), and the skull seems to be of male origin (cranial sexual dimorphism; see [51]). This is so far important, because this subspecies seem to have immigrated to Europe from Asia during the Late Pleistocene, where it is nowaday's extinct [50]. The badger, with its diet (cf. [52]), was not responsible for the bone accumulations of medium-sized mammals and anures, or reptiles, but of micromammals (cf. [53]), also at the Bad Wildungen-Biedensteg open sir site.

Foxes (V lagopus and V! vulpes) might have reused the badger burrows [48]. Foxbones and skulls are typically found at those fox den sites and would explain, additionally, the presence of smaller mammal fox prey remains, especially hares and the micromammal pellets generally found at modern fox dens (cf. [48]).

Quaternary small mustelids in central Europe are rare in the fossil record outside caves (cf. [54, 55]). Their pellets can contain anure or fish bones. Frog or fish remains from Bad Wildungen seem to be partly of prey deposits of Mustela putorius. The small marten type is storing along small rivers or lakes, fishes, frogs, and other animals [48].

A especially high amount of frog bones must have resulted, additionally, from other large water birds and/or other predators which also left pellets and bone remains at the river and along the lake.

7.2. Hyena Population and Cannibalism. The hyena skulls from Bad Wildungen-Biedensteg are from female hyenas which are similar to many other skulls of central Europe (cf. [17]) and are anatomically interesting in their dentition (partly absence of [M.sup.1]), but fall into the variability of C. c. spelaea. A brain case, two incomplete limb bone shafts, a left radius, and a left femur are fitting for a single cub, which are very small in their proportions. They also have bite marks and must have been chewed, as compared to other cannibalistic damaged hyena long bone finds from Europeans caves (cf. [11, 22, 23, 25, 27, 56]). Their proportions fit best for a very young cub, maybe only of a few days or weeks in age, compared to the cub material from the Srbsko-ChlumKomin Cave, Czech Republic [11]. The young hyena was possibly eaten cannibalistically, possibly by another cub, due to competition (cf. modern African hyenas in [57-59]). All bones of the Bad Wildungen hyena population and even the skulls have nibbling, chewing, and cracking marks of hyenas. The lack of the jugals and temporal parts of the skulls is the result of cracking the lower jaws from their joints, which is demonstrated for many skull finds in Europe (cf. [17]). The scavenging of their own species leaves dominantly cranial remains at not only den sites, such as the skulls, lower jaws, and teeth, but also the long bones (e.g., modern spotted hyenas, [60, 61]). Scavenging of their own is best documented in the Srbsko-Chlum-Komin Cave [11]. The dominance of cranial material at Bad Wildungen hyena den site is comparable not only to the German Perick Caves and Rosenbeck Cave and other Sauerland Karst hyena dens, but also to other caves, such as the Czech Sloup Cave, Vypustek, in the Bohemian and Moravian Karst regions [7, 17,56]. Vertebrae and rib bones are underrepresented at most hyena den sites (especially at birthing dens and prey storage den types), the exceptions being where complete articulated skeletons are found at prey storage sites, such as were found at the Czech Vypustek Cave, Koneprusy Cave and SrbskoChlum-Komin Cave [9, 40].

7.3. Hyena Den Type and Recycling of Badger/Fox Dens. Hyena dens are identified starting in the Pliocene to Middle Pleistocene (e.g., [12, 62, 63]). In the Late Pleistocene the hyena den site record is much higher (e.g., [3-6, 8, 17, 64-66]) and more details about the "den type" can be studied. The large bone enrichment at Bad Wildungen was already identified as a product of the activities of C. c.spelaea [35]. The comparison of different Late Pleistocene C. c. spelaea hyena cave and open air den sites in Europe allows a classification of the den type, by separating three main age classes: (1) cubs, (2) adolescents, and (3) adult-senile individuals (Figure 15). The high presence of cubs indicates, similarly as in modern spotted hyenas [57, 67-69], birthing dens. Other indicators for such birthing dens are "nibbling sticks" At Bad Wildungen there are three such chewed bone fragments: one of a mammoth, whose bone fragments are found at birthing dens [70] for teething purposes of hyena cubs [7]; the other nibbling sticks are from Coelodonta and Ursus bone fragments. These birthing dens are generally recycled from medium-sized carnivore, such as porcupines, or by hyenas own excavated burrows, which can be situated nearby commuting dens (cf. modern in [71]). Bad Wildungen must have also been this type of den, where higher amounts of prey remains were accumulated, or even stored (prey storage den type). Similar large bone accumulations at commuting den sites have been reported in Africa from C. c. crocuta (cf. [61, 68, 71-81]).

7.4. Hyena Den Marking. In most cases, pellets of the Late Pleistocene spotted hyenas have repeating shapes, which were found recently at several reported den sites [3, 5-7, 11, 22, 41, 82]. Exact documented excrement markings on a gypsum karst open air den were recently published at the site Westeregeln, Central Germany [9]. A first terminology was published for the pellet shape types [44]. The hyena pellets from Bad Wildungen fall within the hyena pellet shape types. Several smaller pellets are attached to each other, forming spindle-like, or irregular accumulated aggregations, similar to modern African spotted hyena excrements [9]. Modern spotted hyenas are using faecal pellets to mark their territory, especially their den sites [83]. The Ice Age spotted hyenas must have done the same. Well documented examples are found in Germany at two open air sites: Bad Wildungen-Biedensteg [35] and the gypsum karst site Morschen-Konnefeld [84]. Similar abundant pellets are found in caves of France [6] and Czech Republic 5].

7.5. Bone Assemblage and Fauna Statistics. The high amount (10%) of hyena bone remains is typical for Late Pleistocene hyena dens (e.g., [8,11, 65, 66]).

A high percentage of hyena prey bone remains at the site Bad Wildungen-Biedensteg (Figure 16) do not represent the real percentages of the prey. It is more demonstrated, for example, at other hyena open air sites, as a result of taphonomy and selection [9]. The bones of the woolly rhinoceros are extremely massive, and, in contrast to nearly all other large mammal bones, completely filled with the spongiosa. The long bones were difficult or impossible to crack and hyenas always left, in a last stage (stage 3), the bone shaft of long bones or massive bones which are classified in three damage stages [10].

The open air site Bad Wildungen-Biedensteg has delivered only a very few mammoth bones (2% of the prey bones) which are typical at middle high mountainous hyena dens of Europe, where mammoths seem to have been absent or rare [7]. Hyenas specialized there on cave bear scavenging ([42], Figure 16). The amount of Przewalski horse remains (8%) is as usual high. In most open air sites and middle mountainous elevated European caves the small Przewalski horse is the main or second dominant prey (up to 50%; [7, 9-11, 18, 37, 40-44]). If all the small carnivores are excluded from the statistics, then the horse remains represent the second largest prey (cf. [85]). Bones of those horses are recorded with small proportioned forms (see metapod discussion) attributed to E. c. przewalskii in Germany or Czech Republic at other hyena den sites of early to middle Late Pleistocene age [7, 85]. Late Palaeolithic archaeological sites have the youngest records from the Late Magdalenian [86] or Epipalaeolithic/Early Mesolithic [87]. Finally, trackways have been described from the German Volcanic ashes of the Laacher Volcano to be of Przewalski horse origin [37, 88]. Additionally, archaeologists have discussed intensive horse figurations in cave and mobile art and identified also the horses by the unique "M-sign" (resulting from fur colour and fur change) and "uplifted mane" (only in those horses, not in modern present horses) to represent obviously Przewalski horses within the Late Palaeolithic times (cf. e.g., [86, 89]) and especially within the cold periods of the Late Pleistocene.

7.6. Woolly Rhinoceros as Main Prey for Hyenas. Most remains are from the woolly rhinoceros (32%), which corresponds well to several other northern Germany open air hyena den sites, such as Bottrop, Westeregeln, or cave sites on the mountain slope regions, such as Hohle Stein Cave or Teufelskammer Cave ([9, 10, 22, 82], Figure 16). All bones have medium to massive nibbling, chewing, and gnawing marks, mainly produced by the Ice Age spotted hyenas, as compared to other den sites [10, 91] and modern spotted hyenas [92, 93]. Scratches deep into the spongiosa of the joints are very typical of hyena origin and can be found at many other European open air and cave sites (e.g., [11, 21-23, 25, 29, 40, 41, 82, 94, 95]). The material from Bad Wildungen consists of a few cranial and mainly postcranial bones of at least five woolly rhinoceros individuals. Remains of a young, less than one-year-old calf, a young adult female, and a few remains of a male adult skeleton can be distinguished (Figure 5(b)). Besides those, mainly forelimb bones from some other rhinoceros individuals were found. A comparison to a normal bone proportion relation analyses [10] to the material from Bottrop open air site (Figure 5(a)) shows differences mainly in the thoracic (vertebrae, costae) presence. In Bad Wildungen, those thoracic elements are more abundant, similar to those found on nonscavenged skeletons like the Petershagen skeleton [90], which indicates the scavenging of a carcass very nearby the den.

The presence of a carcass is also demonstrated by the articulated vertebral column (Figure 5(b)). To this, most probably, other elements belong. An originally articulated right hind limb (femur and tibia, astragalus, and calcaneus) or forelimb bones, such an ulna and radius, support the original presence of one animal carcass which was decomposed in parts. Such decompositions could have taken days, such as what is known for Late Pleistocene elephant carcasses [43]. The carcass of the most probable female C. antiquitatis must have laid on the right side of her body during main carcass feeding activities, because more bones from that side are preserved. The skull is lacking, but it seems as if all isolated teeth found from the lower jaw indicate the complete destruction of the mandibles by the hyenas. Isolated teeth of woolly rhinoceros are typically at hyena den sites (e.g., [10]). Maybe the skull was cut off by the hyenas or at least destroyed. A few ribs were only cracked, and nearly all are lacking their joints. The long bone joints were not chewed off completely, because of their articulation. This indicates a fresh carcass that was not completely used by the hyenas and was left in an intermediate stage of carcass destruction (cf. Figure 5(a)). After the bone destruction stages, those are in stage 2 sensu Diedrich [10]. The spongiosa remains of woolly rhinoceros were quite often found in the hyena coprolites at the Bad Wildungen-Biedensteg site [35]. The brain case opening of a calf is similarly figured as an adolescent rhinoceros skull from Selm-Ternsche [10], as figured from rhinoceros skull damages from other sites [96].

The finds of juveniles, such as the few-weeks-old rhinoceros (Figures 5(b) and 6), hyena, or the neonate cave bear, fit for the hunting and main activity time of the hyenas at Biedensteg in the late spring and early summer. Other remains of at least four more rhinoceros individuals and other prey remains were imported, possibly from the Ice Age spotted hyenas.

7.7. Hyenas as Cave Bear Scavengers. The cave bear bones might belong to one skeleton of a mature female cave bear [35]. The small diameter, 75 mm, of the scapula glenoid fits for cave bears of the smaller subspecies U. spelaeus subsp. of the early/middle Late Pleistocene, compared, for example, to the cave bear population of the Perick Caves in the Sauerland Karst (Figure 1; [97]) or the newer studied cave bear populations and subspecies of the Rubeland Caves [98]. Also, the other bones and femur fragments were compared to some hundred bones from the Perick and Rubeland Caves, all having again smaller proportions, excluding a U. ingressus cave bear type of the latest Late Pleistocene. Finally, similarly as figured with the "nibbling stick" in the Perick Caves, some cave bear femora and other bone fragment nibbling sticks are present [70], which only hyenas must have produced by teething cubs (cf. [7]). A scavenging of a cave bear carcass outside a cave is the only clear report of such a scenario [97], but is not exceptional, if compared to the hunting/feeding strategies of the Late Pleistocene spotted hyenas. It is now well known that they scavenged cave bear carcasses in the mountain regions of Europe, such as the Sauerland Caves, the Perick Caves, and Rubeland Caves, and additionally several other cave bear dens all over Europe [42, 70, 98, 99].

7.8. Fauna Biodiversity and Climatic Mammoth Steppe Indicators. The faunal statistics demonstrate (Figure 16(b)) that most megafauna bones from Bad Wildungen are related to be of hyena prey origin. Those represent a mammoth steppe megafauna with Coelodonta antiquitatis (cf. [29]), Mammuthus primigenius, Bison/Bos, Megaloceros giganteus, Cervus elaphus, Rangifer tarandus, Equus caballus przewalskii, and boreal mountain forest fauna of Ursus spelaeus subsp. (cf. [35]). Additionally, the pellets include many mammoth steppe environment rodents such as Lemmus lemmus, Dicrostonyx henseli, Microtus gregalis, or Allactaga saliens (cf. [33, 34, 100]). Represented are in higher amounts furthermore birds such as Lagopus lagopus and other species (cf. [34]).

8. Conclusion

The open air hyena den site Bad Wildungen-Biedensteg (NW-Germany) must have been located at the margin of an ancient small lake and the Wilde River in a mammoth steppe landscape on the eastern slopes of the Sauerland Mountains during the early to middle glaciation (early late Pleistocene or Weichselian, about "65.000-90.000 BP," MIS 5c-d). This shallow lake margin, or at least muddy area, was in the center of a large sinkhole structure, which was caused by subsurface dissolution of Zechstein salt in the underground. The sinkhole received freshwater influence by the early Wilde River, indicated by especially freshwater fish remains, but also some other water related animals such as frogs, which were found accumulated in many pellets. Those are excrements of red/arctic foxes, steppe iltis and large carnivore water birds, or owls. Nearby, a badger/fox den burrow area in loess deposits must have been present, where their bone remains and those of their prey (mainly hare, and micromammals) were accumulated, also in pellets. With Biedensteg, an open air hyena birthing and overlapping communal den with prey deposit can be presented with probably reused badger/red fox burrows for the natal den function. 10% of the NISP are Crocuta crocuta spelaea remains, including three grown-up animal skulls, and cranial and postcranial remains of a young cub. Abundant are hyena coprolites (mainly encrusted by caliche), which contain fragments of bones, and most probably quite abundant bone spongiosa fragments from woolly rhinoceros bones. This corresponds to the main hyena prey Coelodonta antiquitatis (NISP = 32%). Another main prey is the horse Equus caballus przewalskii (8%). This dominance of woolly rhinoceros/horses in the Late Pleistocene bone assemblages in northern Europe was caused solely by those large carnivores and is typical of many hyena open air and cave den bone accumulation sites in northern Germany and Czech Republic (central Europe).


The work is dedicated in the memory of L. Lorenz and her husband R. Lorenz (in former times running the Rudolf-Lorenz-Stiftung), who both supported much rescuing material from the locality, research financing, and care keeping of the collection, which is housed in the Heimatmuseum of the City Bad Wildungen. For the kind hosting during the studies the author thanks Mrs. L. Lorenz, and for the cooperation Dr. V. Brendow, the museums head of the Kurmuseum Bad Wildungen. Dr. I. Wrazlido (Leader Museum Natur und Mensch Bielefeld) kindly allowed the comparison of the woolly rhinoceros skeleton from Petershagen. The author thanks Dr. J. Fichter very much for the help in the donation of the collection that was housed partly in the University of Marburg (coll. E. Jacobshagen). The research and project "Hyena open air prey deposit site Bad Wildungen-Biedensteg" was financially supported by the Rudolf-Lorenz-Stiftung, the Sparkassenstiftung of Bad Wildungen, and the PaleoLogic. Sadly, the responsible Ministry of Culture of Hesse and the office for palaeontological monument survey Wiesbaden did not take care of the site, which was damaged further by a newer house construction, where many finds have been destroyed in the late 90s. Finally the author thanks the reviewers, especially Prof. Dr. Muller-Beck, for their supporting critics of the first paper draft. And last, the author would like to thank S. Stevens for the spell check.


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Cajus Diedrich

PaleoLogic Private Research Institute, Petra Bezruce 96, 26751 Zdice, Czech Republic

Correspondence should be addressed to Cajus Diedrich;

Received 15 September 2012; Accepted 27 January 2013

Academic Editor: Atle Nesje

Table 1: Bones of Crocuta crocuta spelaea (Goldfuss 1823) from the
open air site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.       Bone type              Commentary

1       52/45           Skull          Nearly complete, female
2       10at            Skull          Nearly complete, female
3         /             Skull                    --

4       10ev            Skull                Brain case
5       52/51         Mandibula                  --
6         ?           Mandibula                  --

7         ?           Mandibula                  --

8      52/234     Cervical vertebra             Axes
9       10ew           Radius              Without joints
10     52/249     Thoracic vertebra             Disc
11      10em            Femur              Without joints
12     52/209         Coprolite             Single pellet
13     52/220         Coprolite             Single pellet
14     52/207         Coprolite             Single pellet
15     52/210         Coprolite             Single pellet
16     52/212         Coprolite             Single pellet
17     52/206         Coprolite       Single pellet, with prey
                                            bone fragment
18     52/218         Coprolite             Single pellet
19     52/226         Coprolite       Single pellet, with prey
                                            bone fragment
20     52/208         Coprolite       Single pellet, with prey
                                            bone fragment
21     52/225         Coprolite       Single pellet, with prey
                                            bone fragment
22     52/211         Coprolite             Single pellet
23     52/214         Coprolite       Three articulated pellets
24     52/213         Coprolite          Single large pellet
25     52/219         Coprolite            Two articulated
                                            pellets, with
                                         prey bone fragment
26     52/221         Coprolite              Two large
                                         articulated pellets
27     52/237         Coprolite             Single pellet

No.   Left    Right Age    Bite marks         Collection

1            Early adult       x        Rudolf-Lorenz-Stiftung
2            High adult        x        Rudolf-Lorenz-Stiftung
3                 ?                      (Mentioned in [34],
4                Cub           x        Rudolf-Lorenz-Stiftung
5      x     Early adult       x        Rudolf-Lorenz-Stiftung
6                 ?                      (Mentioned in [34],
7                 ?                      (Mentioned in [34],
8               Adult          x        Rudolf-Lorenz-Stiftung
9      x         Cub           x        Rudolf-Lorenz-Stiftung
10               Cub                    Rudolf-Lorenz-Stiftung
11     x         Cub           x        Rudolf-Lorenz-Stiftung
12                                      Rudolf-Lorenz-Stiftung
13                                      Rudolf-Lorenz-Stiftung
14                                      Rudolf-Lorenz-Stiftung
15                                      Rudolf-Lorenz-Stiftung
16                                      Rudolf-Lorenz-Stiftung
17                                      Rudolf-Lorenz-Stiftung

18                                      Rudolf-Lorenz-Stiftung
19                                      Rudolf-Lorenz-Stiftung

20                                      Rudolf-Lorenz-Stiftung

21                                      Rudolf-Lorenz-Stiftung

22                                      Rudolf-Lorenz-Stiftung
23                                      Rudolf-Lorenz-Stiftung
24                                      Rudolf-Lorenz-Stiftung
25                                      Rudolf-Lorenz-Stiftung

26                                      Rudolf-Lorenz-Stiftung

27                                      Rudolf-Lorenz-Stiftung

Table 2: Bones of Ursus spelaeus subsp. Rosenmuller 1794 from the
open air site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.   Bone type       Commentary        Left   Right

1      52/227      Scapula    Without distal part    x
2       52/2       Humerus           Shaft                   x
3      52/241       Ulna          Incomplete         x
4      52/242       Femur          Fragment                  ?

No.    Age    Bite marks         Collection

1     Adult       x        Rudolf-Lorenz-Stiftung
2     Adult       x        Rudolf-Lorenz-Stiftung
3     Adult       x        Rudolf-Lorenz-Stiftung
4     Adult       x        Rudolf-Lorenz-Stiftung

Table 3: Bones of Vulpes vulpes 1758 from the open air prey deposit
site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.      Bone type            Commentary         Left

1       52/39         Cranium             Incomplete
2       52/35         Scapula          Nearly complete
3       52/10         Humerus              Complete
4       52/24         Radius               Complete
5      52/104          Femur           Nearly complete        x
6      52/105a         Tibia               Complete           x
7      52/238        Calcaneus             Complete           x
8      52/127         Pelvis           Nearly complete
9      52/128         Pelvis               Fragment
10     52/239     Metatarsus III           Complete
11     52/240       Metatarsus      Without proximal joint
12      52/21     Lumbar vertebra      Nearly complete
13     52/105b         Costa           Nearly complete

No.   Right   Age Bite marks         Collection

1                 Adult        Rudolf- Lorenz-Stiftung
2       x         Adult        Rudolf-Lorenz-Stiftung
3       x         Adult        Rudolf-Lorenz-Stiftung
4       x         Adult        Rudolf-Lorenz-Stiftung
5                 Adult        Rudolf-Lorenz-Stiftung
6                 Adult        Rudolf-Lorenz-Stiftung
7                 Adult        Rudolf-Lorenz-Stiftung
8                 Adult        Rudolf- Lorenz-Stiftung
9       x         Adult        Rudolf-Lorenz-Stiftung
10      x         Adult         University of Marburg
11                Adult         University of Marburg
12      x         Adult        Rudolf-Lorenz-Stiftung
13      x         Adult        Rudolf-Lorenz-Stiftung

Table 4: Bones of Vulpes lagopus 1758 from the open site Bad
Wildungen-Biedensteg (Hesse, NW-Germany).

No.    Coll.-No.   Bone type                Commentary

1        10bh       Cranium    Nearly complete with right lower jaw
2       52/243     Mandibula             Fragment with P4
3        10eh        Femur                     Shaft
4        10bn       Pelvis             Fragment, acetabulum

No.   left   right    Age     Bite marks          Collection

1                    Senile                Stadtmuseum Bad Wildungen
2      x             Adult                   University of Marburg
3      x                                   Stadtmuseum Bad Wildungen
4      x             Adult                 Stadtmuseum Bad Wildungen

Table 5: Bones of Meles meles Linne 1758 from the open air site Bad
Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.    Bone type          Commentary         Left   Right

1       10ah        Cranium     Skull with lower jaws
2      BadW-1       Cranium     Skull with lower jaws
3       64/1        Humerus         Without joints
4       10ap        Humerus     Without proximal joint    x
5       10ao         Ulna             Incomplete          x
6       10av         Ulna              complete                   x
7       10aw        Radius             Complete           x
8       52/84       Radius          Without joints
9       10ao        Radius             complete                   x
10      52/87       Radius          Without joints
11      10bd       Pisiform            Complete
12      52/86        Femur          Without joints                x
13      52/85        Tibia       Without joints, half     x
14      10aq         Tibia             Fragment
15      10at       Calcaneus           Complete           x
16      10an       Calcaneus           Complete                   x
17      10ay       Astragal            Complete           x
18     BadW-2      Astragal            Complete                   x
19      10qr       Astragal            Complete                   x
20      10lm      Intermedium          Complete
21      10bf      Metatarsus        III, complete         x
22      10bb      Metatarsus         V, complete          x
23     BadW-5     Metatarsus         IV, complete                 x
24     BadW-6     Metatarsus        III, complete                 x
25     BadW-7     Metatarsus         II, complete                 x
26     BadW-8     Metatarsus         I, complete                  x
27     BadW-2     Phalanx II           Complete
28     BadW-3     Phalanx II           Complete
29     BadW-4     Phalanx II           Complete

No.     Age      Bite marks Collection

1      Senile    Rudolf-Lorenz-Stiftung
2     Juvenile   Rudolf-Lorenz-Stiftung
3     Juvenile   Rudolf-Lorenz-Stiftung
4      Adult     Rudolf-Lorenz-Stiftung
5      Adult     Rudolf-Lorenz-Stiftung
6      Adult     Rudolf-Lorenz-Stiftung
7      Adult     Rudolf-Lorenz-Stiftung
8     Juvenile   Rudolf-Lorenz-Stiftung
9      Adult     Rudolf-Lorenz-Stiftung
10    Juvenile   Rudolf-Lorenz-Stiftung
11     Adult     Rudolf-Lorenz-Stiftung
12    Juvenile   Rudolf-Lorenz-Stiftung
13    Juvenile   Rudolf-Lorenz-Stiftung
14     Adult     Rudolf-Lorenz-Stiftung
15     Adult     Rudolf-Lorenz-Stiftung
16     Adult     Rudolf-Lorenz-Stiftung
17     Adult     Rudolf-Lorenz-Stiftung
18     Adult     Rudolf-Lorenz-Stiftung
19     Adult     Rudolf-Lorenz-Stiftung
20     Adult     Rudolf-Lorenz-Stiftung
21     Adult     Rudolf-Lorenz-Stiftung
22     Adult     Rudolf-Lorenz-Stiftung
23     Adult     Rudolf-Lorenz-Stiftung
24     Adult     Rudolf-Lorenz-Stiftung
25     Adult     Rudolf-Lorenz-Stiftung
26     Adult     Rudolf-Lorenz-Stiftung
27     Adult     Rudolf-Lorenz-Stiftung
28     Adult     Rudolf-Lorenz-Stiftung
29     Adult     Rudolf-Lorenz-Stiftung

Table 6: Bones of Mustela putorius Linnaeus 1758 from the open air
prey deposit site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.   Bone type     Commentary      Left   Right    Age

1       10bs       Cranium    Nearly complete                  Senile
2      52/247      Pelvis        Fragment                      Adult

No.   Bite marks          Collection

1                  Stadtmuseum Bad Wildungen
2                    University of Marburg

Table 7: Bones of Mammuthus primigenius (Blumenbach 1799) from the
open air site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.       Bone type               Commentary

1       10ex            Dens             Fragment of lamella
2      52/116     Thoracic vertebra          Neural arch
3      52/149     Thoracic vertebra            Centrum
4      52/222         Long bone       Fragment, "nibbling stick"

No.   Left   Right        Age         Bite marks

1                    Early juvenile
2                       ? Adult           x
3                       ? Adult           x
4      x                 Adult            x

No.         Collection

1     Rudolf-Lorenz-Stiftung
2     Rudolf-Lorenz-Stiftung
3     Rudolf-Lorenz-Stiftung
4     Rudolf-Lorenz-Stiftung

Table 8: Bones of Coelodonta antiquitatis (Blumenbach) from the open
air site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.    Coll.-No.        Bone type             Commentary

1        10ac            Cranium          Middle part with
                                          [M.sup.1] dentition
2        52/37          Mandible         Milk dentition, with
                                         [dm.sub.3], [M.sub.1]
3        52/38          Mandible         Milk dentition, with
                                        [dm.sub.1-3], [M.sub.1]
4        Ma 1             Dens           Milk tooth, upper jaw
5        Ma 2             Dens           Milk tooth, upper jaw
6        Ma 3             Dens           Milk tooth, upper jaw
7        Ma 4             Dens                    P3
8        Ma 5             Dens                    P4
9        Ma 6             Dens                    M1
10       Ma 7             Dens                    M2
11       Ma 8             Dens                    M3
12        10l            Scapula               Fragment
13       52/20           Scapula         Without distal joint
14      52/200           Scapula              Incomplete
15       52/88           Scapula               Fragment
16       180c            Humerus              Incomplete
17        10v            Humerus              Incomplete
18     52/47, 42       Ulna/radius        Shafts, articulated
19    52/116, 111      Ulna/radius        Shafts, articulated
20      52/143            Ulna                   Shaft
21        10p             Ulna                   Shaft
22       52/53            Ulna                   Shaft
23        10a             Ulna                   Shaft
24       52/49           Radius          Without distal joint
25       52/44           Radius                  Shaft
26       52/30           Radius             Proximal joint
27      52/224           Radius              Distal joint
28        10a            Radius             Proximal joint
29      52/235         Intermedium          Nearly complete
30       52/34          Carpale 3           Nearly complete
31       Ma 11        Metacarpale 3         Nearly complete
32       Ma 12        Metacarpale 3         Nearly complete
33      52/101           Phalanx               Complete
34       52/43            Femur                  Shaft
35      52/153            Femur             Shaft, fragment
36       10ab             Femur               Incomplete
37       10ea             Femur                  Shaft
38       10aya            Femur                  Shaft
39      52/228           Patella               Complete
40       52/7             Tibia               Incomplete
41      52/201            Tibia         Without proximal joint
42        10c             Tibia               Incomplete
43       52/9             Tibia         Without proximal joint
44        10t             Tibia         Without proximal joint
45       52/4            Fibula              Distal joint
46       52/16           Fibula                  Shaft
47        10f           Calcaneus             Incomplete
48        10g          Astragalus             Incomplete
49      52/140b        Metatarsus            III, complete
50       Ma 13        Metatarsale 2         Proximal joint
51       Ma 14        Metatarsale 3         Nearly complete
52       Ma 15        Metatarsale 4         Nearly complete
53       52/48           Pelvis               Incomplete
54       52/82           Pelvis               Incomplete
55       52/13           Pelvis             Ilium, fragment
56        10e            Pelvis               Incomplete
57       52/9       Cervical vertebra            Atlas
58       52/1       Cervical vertebra            Axes
59       52/11      Cervical vertebra            No. 3
60       52/18      Cervical vertebra            No. 5
61     52/107-1     Cervical vertebra            No. 6
62     52/107-2     Cervical vertebra            No. 7
63     52/107-3     Thoracic vertebra            No. 1
64        10m       Thoracic vertebra            No. 2
65        10j       Thoracic vertebra            No. 3
66      52/152      Thoracic vertebra       Centrum, No. 4
67     52/108-1     Thoracic vertebra            No. 6
68     52/1808-2    Thoracic vertebra            No. 7
69     52/108-3     Thoracic vertebra            No. 8
70     52/108-4     Thoracic vertebra            No. 9
71        10l       Thoracic vertebra           No. 18
72        10h        Lumbar vertebra             No. 1
73        10r        Lumbar vertebra          Neural arch
74       52/3             Costa                Fragment
75       52/5             Costa                Fragment
76      52/156            Costa                Fragment
77       52/58            Costa         Anterior, 2, distally
78       52/57            Costa         Middle, approx. 6 to 8
79       52/52            Costa           Middle, approx. 4-6
80       52/15            Costa           Middle, approx. 7-9
81      52/100            Costa          Anterior, approx. 2-3
82       52/3a            Costa          Anterior, approx. 3-4
83        10q             Costa          Anterior, approx. 4-6
84        10v             Costa          Anterior, approx. 3-4
85       10ad             Costa                Posterior

No.   Left   Right        Age         Bite marks

1                    Early juvenile       x

2              x     Early juvenile       x

3      x             Early juvenile       x

4                    Early juvenile
5                    Early juvenile
6                    Early juvenile
7              x      Early adult
8              x      Early adult
9              x      Early adult
10     x              Early adult
11     x              Early adult
12     x                ? Adult           x
13     x                 Adult            x
14             x        ? Adult
15                      ? Adult
16             x         Adult            x
17     x                 Adult            x
18     x                Juvenile          x
19             x      Early adult         x
20             x        ? Adult           x
21             x        ? Adult           x
22             x         Adult            x
23             x         Adult            x
24     x                 Adult            x
25             x         Adult            x
26     x                 Adult            x
27             x      Early adult         x
28             x         Adult            x
29             x         Adult
30             x         Adult
31                       Adult
32                       Adult
33                       Adult
34     x                Juvenile          x
35     x
36             x         Adult            x
37     x              Early adult         x
38     x              Early adult         x
39             x         Adult
40             x         Adult            x
41             x         Adult            x
42     x                 Adult            x
43             x         Adult            x
44             x         Adult            x
45     x                 Adult            x
46     x                 Adult            x
47             x         Adult            x
48             x         Adult            x
49             x         Adult
50                       Adult
51                       Adult
52                       Adult
53     x                 Adult            x
54             x         Adult            x
55     x                 Adult            x
56     x                 Adult            x
57                    Early adult         x
58                    Early adult         x
59                    Early adult         x
60                    Early adult
61                    Early adult         x
62                    Early adult         x
63                    Early adult         x
64                    Early adult         x
65                    Early adult         x
66                    Early adult         x
67                    Early adult         x
68                    Early adult         x
69                    Early adult         x
70                    Early adult         x
71                    Early adult         x
72                    Early adult         x
73                    Early adult         x
74                         ?              x
75                         ?
76                    Early adult
77     x              Early adult

78     x              Early adult         x
79             x      Early adult         x
80     x              Early adult         x
81             x      Early adult         x
82             x      Early adult         x
83     x              Early adult         x
84     x              Early adult         x
85             x      Early adult         x

No.            Collection

1        Rudolf-Lorenz-Stiftung

2        Rudolf-Lorenz-Stiftung

3        Rudolf-Lorenz-Stiftung

4         University of Marburg
5         University of Marburg
6         University of Marburg
7         University of Marburg
8         University of Marburg
9         University of Marburg
10        University of Marburg
11        University of Marburg
12       Rudolf-Lorenz-Stiftung
13       Rudolf-Lorenz-Stiftung
14       Rudolf-Lorenz-Stiftung
15       Rudolf-Lorenz-Stiftung
16    (Mentioned in [34], missing)
17       Rudolf-Lorenz-Stiftung
18       Rudolf-Lorenz-Stiftung
19       Rudolf-Lorenz-Stiftung
20       Rudolf-Lorenz-Stiftung
21       Rudolf-Lorenz-Stiftung
22       Rudolf-Lorenz-Stiftung
23       Rudolf-Lorenz-Stiftung
24       Rudolf-Lorenz-Stiftung
25       Rudolf-Lorenz-Stiftung
26       Rudolf-Lorenz-Stiftung
27       Rudolf-Lorenz-Stiftung
28       Rudolf-Lorenz-Stiftung
29       Rudolf-Lorenz-Stiftung
30       Rudolf-Lorenz-Stiftung
31        University of Marburg
32        University of Marburg
33       Rudolf-Lorenz-Stiftung
34       Rudolf-Lorenz-Stiftung
35       Rudolf-Lorenz-Stiftung
36       Rudolf-Lorenz-Stiftung
37       Rudolf-Lorenz-Stiftung
38       Rudolf-Lorenz-Stiftung
39       Rudolf-Lorenz-Stiftung
40       Rudolf-Lorenz-Stiftung
41       Rudolf-Lorenz-Stiftung
42       Rudolf-Lorenz-Stiftung
43       Rudolf-Lorenz-Stiftung
44       Rudolf-Lorenz-Stiftung
45       Rudolf-Lorenz-Stiftung
46       Rudolf-Lorenz-Stiftung
47       Rudolf-Lorenz-Stiftung
48       Rudolf-Lorenz-Stiftung
49       Rudolf-Lorenz-Stiftung
50        University of Marburg
51        University of Marburg
52        University of Marburg
53       Rudolf-Lorenz-Stiftung
54       Rudolf-Lorenz-Stiftung
55       Rudolf-Lorenz-Stiftung
56       Rudolf-Lorenz-Stiftung
57       Rudolf-Lorenz-Stiftung
58       Rudolf-Lorenz-Stiftung
59       Rudolf-Lorenz-Stiftung
60       Rudolf-Lorenz-Stiftung
61       Rudolf-Lorenz-Stiftung
62       Rudolf-Lorenz-Stiftung
63       Rudolf-Lorenz-Stiftung
64       Rudolf-Lorenz-Stiftung
65       Rudolf-Lorenz-Stiftung
66       Rudolf-Lorenz-Stiftung
67       Rudolf-Lorenz-Stiftung
68       Rudolf-Lorenz-Stiftung
69       Rudolf-Lorenz-Stiftung
70       Rudolf-Lorenz-Stiftung
71       Rudolf-Lorenz-Stiftung
72       Rudolf-Lorenz-Stiftung
73       Rudolf-Lorenz-Stiftung
74       Rudolf-Lorenz-Stiftung
75       Rudolf-Lorenz-Stiftung
76       Rudolf-Lorenz-Stiftung
77       Rudolf-Lorenz-Stiftung

78       Rudolf-Lorenz-Stiftung
79       Rudolf-Lorenz-Stiftung
80       Rudolf-Lorenz-Stiftung
81       Rudolf-Lorenz-Stiftung
82       Rudolf-Lorenz-Stiftung
83       Rudolf-Lorenz-Stiftung
84       Rudolf-Lorenz-Stiftung
85       Rudolf-Lorenz-Stiftung

Table 9: Bones of Bison priscus (Bojanus 1827) from the open air site
Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.       Bone type                  Commentary

1         /             Dens                   M1, upper jaw
2         /             Dens                   M1, lower jaw
3         /            Scapula
4      BadW-9          Scapula                 Proximal half
5       10af         Metacarpus                Proximal joint
6         /         Carpale 3 + 4
7      52/205           Femur         Distal joint and shaft fragment
8        10o            Femur                      Shaft
9        10k            Femur              Distal joint, fragment
10     52/236           Tibia              Without proximal joint
11      52/12         Calcaneus               Nearly complete
12      52/17     Thoracic vertebra               Centrum

No.   Left   Right    Age    Bite marks

4                    Adult
5      x             Adult       x
7      x             Adult       x
8              x     Adult       x
9              x     Adult       x
10             x     Adult       x
11             x     Adult       x
12                   Adult       x

No.                    Collection

1             (Mentioned in [34], missing)
2             (Mentioned in [34], missing)
3             (Mentioned in [34], missing)
4                Rudolf-Lorenz-Stiftung
5                Rudolf-Lorenz-Stiftung
6             (Mentioned in [34], missing)
7                Rudolf-Lorenz-Stiftung
8                Rudolf-Lorenz-Stiftung
9              Stadtmuseum Bad Wildungen
10    Museum Korbach, (Stadtmuseum Bad Wildungen)
11               Rudolf-Lorenz-Stiftung
12               Rudolf-Lorenz-Stiftung

Table 10: Bone material list of Equus caballus przewalskii
Poljakoff 1881 from the open air prey deposit site Bad
Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.        Bone type               Commentary

1      52/221          Mandibula             Nearly complete
2       52/27          Mandibula           Anterior part, male
3      52/203           Cranium            Occipital, fragment
4      52/147             Dens                   C, male
5       52/50         Ulna/radius              Incomplete
6       10aa          Ulna/radius            Nearly complete
7      52/112       length = 236 mm,         Nearly complete
                  distal width = 50 mm
8      52/155          Metacarpus             Distal joint
9       52/14          Phalanx 1                Complete
10      52/78          Phalanx 2                Complete
11      10lt        length = 257 mm,            Complete
                  distal width = 53 mm
12      52/51            Tibia                  Fragment
13      52/28            Pelvis              Fragment, ilium
14       10i             Pelvis              Fragment, ilium
15     52/131      Cervical vertebra      Fragment, neural arch
16     52/202      Cervical vertebra      Fragment, neural arch
17      10eq        Lumbar vertebra      No. 4, without processi
18      10ad             Pelvis            Sacrum, incomplete
19     52/157            Costa                  Fragment

No.   Left   Right     Age      Bite marks         Collection

1      x              Adult                  Rudolf-Lorenz-Stiftung
2                    Juvenile                Rudolf-Lorenz-Stiftung
3                                            Rudolf-Lorenz-Stiftung
4                     Adult                  Rudolf-Lorenz-Stiftung
5              x      Adult         x        Rudolf-Lorenz-Stiftung
6              x      Adult         x        Rudolf-Lorenz-Stiftung

7      x              Adult         x        Rudolf-Lorenz-Stiftung

8      x              Adult                  Rudolf-Lorenz-Stiftung
9                                            Rudolf-Lorenz-Stiftung
10                                           Rudolf-Lorenz-Stiftung

11             x      Adult         x        Rudolf-Lorenz-Stiftung

12                                  x        Rudolf-Lorenz-Stiftung
13     x              Adult         x        Rudolf-Lorenz-Stiftung
14             x      Adult         x        Rudolf-Lorenz-Stiftung
15                    Adult                  Rudolf-Lorenz-Stiftung
16                    Adult         x        Rudolf-Lorenz-Stiftung
17                   Juvenile       x        Rudolf-Lorenz-Stiftung
18                   Juvenile       x        Rudolf-Lorenz-Stiftung
19                                  x        Rudolf-Lorenz-Stiftung

Table 11: Bones of Megaloceros giganteus (Blumenbach 1799) from the
open air site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.       Bone type           Commentary       Left

1         /           Mandibula       Fragment with M1-3
2         /             Dens            P1, upper jaw
3         /             Dens            M2, upper jaw       x
4         /             Dens            M3, upper jaw       x
5         /       Cervical vertebra         Atlas
6         /       Cervical vertebra          Axes
7       52/32           Tibia            Distal joint

No.   Right    Age    Bite marks            Collection

1       x                          (Mentioned in [34], missing)
2                                  (Mentioned in [34], missing)
3                                  (Mentioned in [34], missing)
4                                  (Mentioned in [34], missing)
5                                  (Mentioned in [34], missing)
6                                  (Mentioned in [34], missing)
7       x     Adult       x           Rudolf-Lorenz-Stiftung

Table 12: Bones of Rangifer tarandus Linne 1758 from the open air
site Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.    Bone type                Commentary

1         /          Dens                       --
2         /          Dens                       --
3       52/40       Antler      Dropped antler with base, fragment
4       52/41       Antler      Dropped antler with base, fragment
5       52/33       Antler      Dropped antler with base, fragment
6      52/132       Scapula                 Incomplete
7      52/126       Scapula                 Incomplete
8     52/115-1       Ulna                 Proximal joint
9     52/115-3      Radius                 Distal joint
10    52/115-4      Radiale                  Complete
11    52/115-5    Intermedium                Complete
12    52/115-8      Carpale                  Complete
13    52/115-7     Carpale 4                 Complete
14     52/117     Metacarpus               Distal joint
15      52/52       Pelvis             Acetabulum, fragment
16      52/57       Pelvis             Acetabulum, fragment
17    52/115-2     Phalanx 1     Without proximal joint, forelimb
18    52/115-6     Phalanx 2         Proximal joint, forelimb
19      52/74        Tibia               Fragment, distal
20     52/151        Tibia                Nearly complete
21      52/10        Tibia            Without proximal joint
22      10lz       Phalanx 1     Without proximal joint, hind limb
23     52/246      Phalanx 1     Without proximal joint, hind limb
24     4.4/54      Phalanx 1                 Complete

No.   Left   Right     Age      Bite marks

3      x              Adult         x
4              x      Adult         x
5      x              Adult         x
6              x      Adult         x
7      x              Adult         x
8                    Juvenile
9              x     Juvenile
10             x     Juvenile
11             x     Juvenile
12             x     Juvenile
13             x     Juvenile
14                   Juvenile
15                    Adult
16                    Adult
17                   Juvenile
18                   Juvenile
19                   Juvenile
20             x      Adult         x
21     x              Adult         x
22                   Juvenile
23                   Juvenile
24                    Adult

No.            Collection

1     (Mentioned in [34], missing)
2     (Mentioned in [34], missing)
3        Rudolf-Lorenz-Stiftung
4        Rudolf-Lorenz-Stiftung
5        Rudolf-Lorenz-Stiftung
6        Rudolf-Lorenz-Stiftung
7        Rudolf-Lorenz-Stiftung
8        Rudolf-Lorenz-Stiftung
9        Rudolf-Lorenz-Stiftung
10       Rudolf-Lorenz-Stiftung
11       Rudolf-Lorenz-Stiftung
12       Rudolf-Lorenz-Stiftung
13       Rudolf-Lorenz-Stiftung
14       Rudolf-Lorenz-Stiftung
15       Rudolf-Lorenz-Stiftung
16       Rudolf-Lorenz-Stiftung
17       Rudolf-Lorenz-Stiftung
18       Rudolf-Lorenz-Stiftung
19       Rudolf-Lorenz-Stiftung
20       Rudolf-Lorenz-Stiftung
21       Rudolf-Lorenz-Stiftung
22       Rudolf-Lorenz-Stiftung
23       Rudolf-Lorenz-Stiftung
24       Rudolf-Lorenz-Stiftung

Table 13: Bones of Cervus elaphus Linne 1758 from the open air site
Bad Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.   Bone type         Commentary        Left   Right

1       10ep       Cranium     Maxillar, with M1-2             x
2     52-113-1    Metatarsus   Without distal joint            x

No.     Age      Bite marks         Collection

1     Juvenile                Rudolf-Lorenz-Stiftung
2     Juvenile                Rudolf-Lorenz-Stiftung

Table 14: Bones of Lepus sp. from the open air site Bad
Wildungen-Biedensteg (Hesse, NW-Germany).

No.   Coll.-No.           Bone type

1        98                Cranium

2        14                Cranium
3        63h               Cranium
4        12               Mandibula
5        11               Mandibula
6        63g               Humerus
7        63f               Radius
8        15              Radius/ulna
9        63b                Femur

10       63c                Femur

11       10                 Femur
12       63d                Tibia

13        9                 Tibia
14       13                 Tibia
15       63e                 Pes

16     52-105c            Calcaneus
17        3                Pelvis
18        5                Pelvis
19      52/10               Femur
20     52/248          Lumbar vertebra
21       63a      Pelvis and lumbalvertebra
22     52/249             Calcaneus
23     52/252                Pes
24     52/253           Metatarsus IV
25     52/254            Astragalus
26     52/251            Metacarpus
27     52/256               Ulnar
28     52/255             Tarsalia

No.             Commentary             Left   Right     Age

1         Brain case, frontals,                        Adult
          parietals, incomplete
2                Maxillar               x              Adult
3                Maxillar                       x      Adult
4               Incomplete              x              Adult
5               Incomplete                      x     Juvenile
6          Half, from skeleton                  x      Adult
7          Half, from skeleton                  x      Adult
8             Without joints                    x      Adult
9       Distal joint incomplete,                x      Adult
              from skeleton
10     Half without distal joint,       x              Adult
              from skeleton
11            Without joints            x             Juvenile
12     Proximal joint incomplete,               x      Adult
              from skeleton
13        Without proximal joint                x     Juvenile
14         Without middle shaft                 x      Adult
15    Nearly complete articulated,              x      Adult
              from skeleton
16               Complete                       x      Adult
17         Fragment, acetabulum         x              Adult
18         Fragment, acetabulum         x              Adult
19              Incomplete              x             Juvenile
20              Incomplete                            Juvenile
21      Articulated from skeleton                      Adult
22              Incomplete                      x     Juvenile
23       Incomplete, articulated        x              Adult
24               Complete               x              Adult
25               Complete                       x      Adult
26             3 incomplete                           Juvenile
27              2 complete              x       x      Adult
28              2 complete                      x      Adult

No.   Bite marks         Collection

1                    University Marburg

2                    University Marburg
3                  Rudolf-Lorenz-Stiftung
4                    University Marburg
5                    University Marburg
6                  Rudolf-Lorenz-Stiftung
7                  Rudolf-Lorenz-Stiftung
8         x          University Marburg
9                  Rudolf-Lorenz-Stiftung

10        x        Rudolf-Lorenz-Stiftung

11                   University Marburg
12                 Rudolf-Lorenz-Stiftung

13                   University Marburg
14        x          University Marburg
15                 Rudolf-Lorenz-Stiftung

16                 Rudolf-Lorenz-Stiftung
17                   University Marburg
18                   University Marburg
19                   University Marburg
20                   University Marburg
21                 Rudolf-Lorenz-Stiftung
22                 Rudolf-Lorenz-Stiftung
23                 Rudolf-Lorenz-Stiftung
24                 Rudolf-Lorenz-Stiftung
25                 Rudolf-Lorenz-Stiftung
26                 Rudolf-Lorenz-Stiftung
27                 Rudolf-Lorenz-Stiftung
28                 Rudolf-Lorenz-Stiftung
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Author:Diedrich, Cajus
Publication:Journal of Geological Research
Article Type:Report
Geographic Code:4EUGE
Date:Jan 1, 2013
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