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Quaternary International 436 (2017) 76e83

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Quaternary International

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Homotherium from Middle archaeological and den sites of Germany e , taphonomy and a revision of the Schoningen,€ West Runton and other saber-tooth sites

* Cajus G. Diedrich a, b, , 1, Donald A. McFarlane c a PaleoLogic, Private Research Institute, Petra Bezruce 96, CZ-26751 Zdice, Czech Republic b University of Koblenz-Landau, IfIN, Department of Biology, Universitatsstr.€ 1, D-56070 Koblenz, Germany c Keck Science Department, The Claremont Colleges, 925 North Mills Avenue, Claremont, CA 91711-5916, USA article info abstract

Article history: Four new saber-tooth cat () sites in Germany with new dental and postcranial bone ma- Received 19 April 2016 terial are different in their taphonomic context: 1. The Archaeological Middle Palaeolithic (MIS 9e- Received in revised form Interglacial) Schoningen€ Lake site with remains of a cub carcass, 2. The Middle Palaeolithic (MIS 5e-9) 7 October 2016 Archaeological/ den site of Cave yielding a lower of an older individual, 3. Accepted 13 October 2016 The Zoolithen Cave (MIS 3e9) cave bear/ den with one distal half humerus of an adult, 4. The Available online 1 March 2017 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. Keywords: e Homotherium 330.000 315.000 BP), all other material seems to come from the same Middle Pleistocene warm period, Holsteinian Interglacial (MIS 9e) or few younger Saalian interstadials (MIS 7a, e) deposits, and did not extend over the last MIS 7 glacial Germany into the . Homotherium as hyena-like slow moving cat seems to have disappeared within Archaeological camp versus hyena den the Saalian due to competition with other like Ice Age spotted and brown (Crocuta context crocuta praespelaea/ultima and Pachycrocuta brunnea mosbachensis). The juvenile saber-tooth cat cub and Cuon bite damage from Schoningen€ might be in archaeological context or represent only a carnivore kill. At the Zoolithen versus saber tooth cat humeri Cave, the single bone must have been imported into a hyena prey bone assemblage. The situation is No human tool possibly similar at the two other sites Ketsch and . 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 -developed supracondylar ridge distinguishes Homotherium well from Middle/Late Pleistocene 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 P4/M1 impact marks around the trochlea. 1e2 mm small, mostly triangular-oval bite marks on the lion humerus shaft compacta results from a second and not from “Neanderthal tool use”. Those bite mark sizes are produced mainly of the upper teeth of a the red Cuon alpinus subsp. (or small fox Vulpes praecorsac), which were present in the region within the Holsteinian/Saalian. © 2016 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction monograph by Ballesio (1963). This specimen includes complete preserved humeri, which are compared herein for a revision of The only known European saber-tooth cat Homotherium cren- several questionable “distal humeri” described in recent years from atidens (Weithofer, 1889) skeleton from the Early Pleistocene (Vil- several Central European sites often as “felid or lion humeri” (e.g. lafranchian) of the French site Seneze is well published in a Lewis et al., 2010). The Middle Pleistocene European saber-tooth , which are not yet known by skeletons, are assigned to Homotherium latidens (Owen, 1843) (e.g. Anton et al., 2005), for * Corresponding author. PaleoLogic, Private Research Institute, Petra Bezruce 96, which one of the most northern records comes from an English CZ-26751 Zdice, Czech Republic. cave (Dawkins and Sandford, 1872; McFarlane and Lundberg, 2013). E-mail addresses: [email protected] (C.G. Diedrich), DMcFarlane@kecksci. “ ” claremont.edu (D.A. McFarlane). The Late Pleistocene records from the North Sea (Reumer et al., 1 www.paleologic.eu. 2003) are without stratigraphic value or reliable dates because of http://dx.doi.org/10.1016/j.quaint.2016.10.015 1040-6182/© 2016 Elsevier Ltd and INQUA. All rights reserved. C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 77 their embedding in modern marine sediments: no other European hyena feeding strategies on their felid competitors, lions and saber- finds support their survival into the last Ice Age. tooth cats. The material found in Germany (Fig. 1A) is quite sparse and lacks even complete skulls. A few single teeth and postcranial elements 2. Material and methods have been published. The Early Pleistocene records are from the river terrace site Untermabfeld (cf. Hemmer et al., 2011). Some The carnivore material of the Paleon - Schoningen€ Spear material comes from the Mosbach Sands Neckar River terrace Museum (¼ Pal) was only screened and is partly exposed in the (Schütt, 1970), which are revised by the actual stratigraphy of this permanent exhibition, which needs further detailed work in future. site (cf. Stephan, 2014) now into the early Middle Pleistocene The humerus discussed herein was checked by the tooth mark Cromerian Complex. Further Middle Pleistocene material was forms and sizes. The studied saber-tooth cat and other bone ma- found in the “Heppenloch” also being called Gutenberg Cave terial of the Balve Cave is stored in the Geologisch-Palaontologische€ (Adam, 1975). Museum of the Westphalian Wilhelms-University Münster (¼ In this contribution, Middle Pleistocene material from three new GPIM). The Zoolithen Cave material is in the collection of the German sites can be added (Figs. 1e3). It is not intended to be a full Frankische€ Hohlen€ und Karstforscher Club (¼ FHKF). One humerus review of German, or revision of Eurasian Pleistocene saber-tooth fragment is stored in the Staatliche Museum für Naturkunde cat sites, but it is of urgent need because material has been Stuttgart (¼ SMNS). One complete lion humerus is stored in the confused. Some saber-tooth cat remains are figured (only 0,1%, in Landesamt Sachsen Anhalt collection (¼ LDA). The upper jaw saber- each publication) in lion publications (cf. Diedrich, 2011a,b; tooth cat canine from the Czech C718 Cave site is in the Natio- Diedrich and Rathgeber, 2012). Opposite, some lion remains have nalmuseum Prague (¼ NMP). The compared damaged lion hu- been misidentified as saber-tooth cat remains. For example, the merus from the Neander Valley is in the Goldfuss-Museum of the newest “saber-tooth cat” records described by Serangeli et al. Rheinische Wilhelms-University Bonn (¼ GMB). A single tooth cusp (2015) represent only one saber-tooth cat cub dentition and to its from Sanderhausen is kept in the collection of the Spenglermu- age class fitting humerus such as few other bone fragments of a cub seum Sangerhausen (¼ SPM). (not two individuals as discussed therein). We revise the figured humerus determined in their study as “Homotherium” (second in- dividual) here. Furthermore, the well-known taphonomy of Euro- 2.1. Geology and dating pean Late Pleistocene lion carcasses caused mainly by hyena feeding (Diedrich, 2011a) can be extended with more details of All new German sites are of Middle Pleistocene (Cromerian- Saalian, ¼ MIS 21-7) in age according to the updated German

Fig. 1. Pleistocene saber-tooth cat Homotherium sites in Germany and Czech Republic and other compared sites in England (dating composed from Fejfar, 1956; Schütt, 1970; Adam, 1975; Stuart and Lister, 2010; Serangeli et al., 2015; with new German sites being added). 78 C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83

Fig. 2. Early-Middle Pleistocene saber-tooth cat Homotherium remains from German and Czech sites (see Fig. 1). 1. H. crenatidens, left upper jaw canine of an adult saber-tooth cat from the Konĕprusy Doline, Bohemian , Czech Republic (coll. NMP). 2. H. latidens, left lower jaw canine of an older cat from the Balve Cave, Karst, Germany (modified from Diedrich, 2011a) (coll. GPIM). 3. Upper C fragment from Sanderhausen (D), lingual (coll. SPM). 4e8. H. latidens, incomplete upper and lower jaw tooth set (with well serrated cutting edges and non-closed roots) of an older cub (half to maximum one year) from Schoningen€ Lake shore site, Germany (modified from Serangeli et al., 2015): 5. Canine 3 4 fragment. Right I , 6. Left P , 7. Right lower C, 8. Right M1.9.H. latidens, coxa from the Zoolithen Cave, Upper Franconia Karst, Germany (modified from Diedrich, 2011d), (coll. FHKF), lateral. stratigraphic chart (cf. Stephan, 2014;cf.Fig. 1B). The H. latidens re- which the Holsteinian Interglacial is dated into the MIS 9e isotope mains from Schoningen€ are dated according to the newest work on stage (330.000e315.00 BP) (Müller, 1974). It is characterized by the the Middle to Late Pleistocene stratigraphy in northern Germany, in development in general of temperate mixed forests in central and

Fig. 3. Early-Middle Pleistocene saber-tooth cat Homotherium humeri from European sites. 1. Humerus (mirrored) from the H. crenatidens (Weithofer, 1889) skeleton of Seneze (F) (from Ballesio, 1963), cranial. 2. Humerus from Incarcal (Es) (from Anton et al., 2005), cranial. 3. Distal humerus from C718 Cave (CZ) (coll. NMP), caudal. 4. Right distal humerus from West Runton (GB) (from Lewis et al., 2010). 5. Left distal half humerus of a grown up from the Zoolithen Cave (D), cranial and mirrored (modified from Diedrich, 2011d). 6. Left distal half humerus of a grown up cat from the Rhine River Terrace Ketsch-Kreuzwiese (coll. SMNS), cranial. C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 79 northwestern and seasonality (e.g., Nitychoruk et al., 2005; H. crenatidens and Middle Pleistocene H. latidens (Figs. 1 and Koutsodendris et al., 2010). The Zoolithen Cave has been recently 3). The humerus differences of both biostratigraphic relevant spe- revised and its bone remains are thought to have been accumulated cies seem not to relate to or variability in sizes, not only in Late Pleistocene times, but instead since the Elsterian/ as thought to have been demonstrated for saber-tooth cat Holsteinian (Diedrich, 2014). The long-term multiple use of this cave of Europe and Northern America (Anton et al., 2014). As as a cave bear, hyena and wolf den within different climatic periods a simple fact of a strong faunal turnover between the Early/Mid- explains the large amount of bones (mainly different cave bear spe- Pleistocene, it must be expected, that there was also a species cies/subspecies of Middle Pleistocene: Ursus deningeri,LatePleisto- change/evolutionary trend within Homotherium, similar as for cene: Ursus spelaeus eremus/spelaeus, Ursus ingressus) and the other felids (e.g. Hemmer, 2004). presence of the “largest amount of lion remains ever excavated in a The Early Pleistocene holotype skeleton of H. crenatidens is European cave”, which also includes Middle Pleistocene P. l. fossilis recently under new studies, and original material was not possible remains (Diedrich, 2011a, 2014). All material, including the rare include. The most obvious character to distinguish the humeri of saber-tooth remains, are in secondary/tertiary positions within the H. crenatidens and H. latidens lies in the difference of the complete, cave system due to predator activities and later floods and can not be and especially the distal shaft width, which is about 5 mm larger dated exactly in many cases, except using biostratigraphy (e.g. and more robust in the older H. crenatidens (Fig. 3). Based on the C. alpinus spelaeus and H. latidens are such as U. deningeri Middle Early Pleistocene Seneze skeleton H. crenatidens humerus, the Early Pleistocene species: Diedrich, 2014). The third site discussed here, Pleistocene Homotherium humerus figured from the Spanish site Ketsch, is a Rhine River Valley gravel pit terrace site, which has mixed Incarcal described as “H. latidens” (cf. Galobart et al., 2003; Anton Middle to mainly Late elements (Diedrich and et al., 2005) must be revised herein to belong to H. crenatidens, Rathgeber, 2012). also correlating then with the biostratigraphy (Fig. 1B). Also Italian finds (cf. Sardella and Iurino, 2012) seem to belong to this older 3. Results and discussion species. The humeri from the Middle Pleistocene H. latidens are slimmer 3.1. Saber-tooth cat Homotherium versus lion Panthera humerus or gracile in their shafts in all herein figured English and German osteology Middle Pleistocene specimens (Fig. 2). The comparison between the humeri figured herein (Fig. 3) includes the “P. leo” humerus from In most previous accounts of Middle Pleistocene H. latidens,the the Cromerian West Runton interglacial MIS 19 (Stuart and Lister, distal humeri were figured, but in no case was the most important 2010) site in England (cf. Lewis et al., 2010). This humerus is, defining character discussed, although it was also used in soft tissue compared to the H. crenatidens holotype humerus in same size and fore limb reconstructions by Anton et al. (2009). Based on the com- shape in the distal shaft width to the German finds (Fig. 2). The plete and well-preserved Plio-/Early Pleistocene H. crenatidens hu- English West Runton site also delivered a calcaneus of H. latidens merus figured by Ballesio (1963) (Fig. 3.1), the supracondylar ridge is (Lewis et al., 2010), which finally supports the humerus absent in this , which does not allow a rotation of the paw (cf. identification. Anton et al., 2009). Here, in one example figured (Fig. 4A), for a small lion (most probaly female) from the Late Pleistocene of the Her- 3.3. Hyena bone damage versus human “bone tools” mann's Cave (Harz Mountain Range), the ridge is strong and about 2e3mmwidee a level of development similar to that seen in The taphonomy of carnivore bone damage for the Schoningen€ modern and Middle to Late Pleistocene lions (P. l. leo, P. l. spelaea, P. l. site is misinterpreted in both papers of Berkholst (2015) and fossilis:cf.Dietrich,1968; Diedrich, 2011a,b). On this ridge, the rotator Serangeli et al. (2015). Furthermore, the were not yet musculature (Brachiradialis) for the lower forelimb is attached to the well determined (c.f Kolfschoten, 2014), but are well known now in radius (e.g. Frewein and Vollmerhaus, 1994; Anton et al., 2009) most cases (see Table 1). Older literature and important material (Fig. 4A). This allows lions to catch its prey with the forelimb paws has been overlooked, for example the only complete stripped hyena (Anton et al., 2009). It is much different in the saber-tooth cat ancestor skull of Parahyaena brunnea mosbachensis (cf. Reichenau, Homotherium (as in hyenids, canids), where absence of the supra- 1906; Schütt, 1971), to which material another partial skull can condylar ridge does not allow the forelimbs to be rotated and used in be added from the Middle Pleistocene of the Upper Rhine Graben prey capture (Anton et al., 2009). Also, based on this osteological herein as very rare record in Central Europe (Fig. 3). The spotted character, it is likely that the gait of Homotherium was “hyena e like” and brown hyenas are still to be discussed, if those are late Early (Anton et al., 2005, 2009, 2014). The supracondylar ridge is also ab- Pleistocene Parahyaena brunnea mosbachensis (in some cases sent in the two distal humeri from the Zoolithen Cave and Ketsch Pachycrocuta: e.g. Palmqvist et al., 2011)orCrocuta crocuta praes- river terrace site (Fig. 3). Using this very important ostelologic char- pelaea/ultima (cf. Schütt, 1971; Tseng and Chang, 2007) forms in the acter, the felid humerus from Schoningen€ figured by Serangeli et al. Mid-Pleistocene. Whereas Berkholst (2015) identified bite damage (2015) (Fig. 4A) is a typical lion humerus by comparison to the on a distal bovid tibia as “rodent bite marks”, those should have H. crenatidens holotype humerus (cf. Ballesio, 1963: Fig. 3). Interest- been compared to the larger rodent bite marks of species such as ingly, the length of the supracondylar ridge is 5e10 mm shorter in the Pleistocene porcupines, which are completely different (cf. Schoningen€ specimen than in other Middle-Late Pleistocene lions, is Diedrich, 2009). The bite marks on this bovid guilt bone (see some such as the Late Pleistocene Hermann's Cave lion humerus Fig. 4B) are in fact typical large predator bite marks, based on bite (Fig. 4A). The Mid-/Late Pleistocene lions from Germany are still in mark analysis of large prey (e.g elephants, and even revision about this theme of “three different sized” forms (not vari- : Diedrich, 2005, 2012, 2013b). Instead of classical “bite mark ability nor sexual dimorphism) are not discussed herein further. metric” analyses, the much more successful direct digital- projection test of the “bite-fit” into the bite marks of different 3.2. The humeri osteologic and biostratigraphic differences in predator candidates was applied to the lion humerus and the bovid H. crenatidens/latidens tibia from Schoningen€ (Fig. 4B). The test of the largest predators Homotherium, Panthera, Hyeana and Crocuta dentitions demon- The material from Germany is few and most is incomplete, but strates that in both cases only hyenas account for all the bite the humeri allow a new discussion to the validities of the Early damage (see Fig. 4). Most convincing is the fit of the cutting scissor 80 C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83

Fig. 4. A. Comparison of small-sized Middle-Late Pleistocene lion humeri (most probably females) and their taphonomic damage stages caused by different hyeana species. The “non-Homotherium” distal humerus (modified from Serangeli et al., 2015) from the archaeological site Schoningen€ lake in comparison to similar sized lion humeri from Late Pleistocene German (1. Hermann's Cave cave bear den, 2. Teufelskammer den). The typical lion character (Supracondylar ridge) and also much thicker and not oval shaft (cf. differences in Homotherium humeri Fig. 3) let identify this humerus fragment as a lion subspecies. Furthermore, the taphonomic felid humeri comparison demonstrate the chew/cut damage stages made by hyenas e most probably the Middle Pleistocene Parahyaena brunnea mosbachensis (new unpublished record added herein from the Rhine Graben, C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 81

Table 1 Updated megafauna list of the Middle Pleistocene Schoningen€ Pre-Neanderthal hunter camp and Homotherium lake shore site with “mixed forest/grasland” palaeoenvironmnet (compiled from Kolfschoten, 2014; Berkholst, 2015; Serangeli et al., 2015; and herein added or corrected), and comparison to the Middle Pleistocene nearby (about 30 km) Bauman's Cave with mountain forest fauna (from Diedrich, 2013a,c).

Schoningen€ Lake shore Baumann's Cave Harz Foreland (MIS 9e Interglacial) Harz Mountain Range (MIS 6e9 Stadials/Interstadials)

Felidae Panthera leo subsp. (small lion) Panthera leo subsp. (small lion) (added herein) Homotherium latidens (corrected herein) Panthera leo fossilis (large lion) Hyaenidae Hyaenidae Parahyeana brunnea cf. mosbachensis sp. (added herein) Crocuta crocuta praespelaea/ultima Canidae Canidae Cuon alpinus spelaeus (added herein) Cuon alpinus spelaeus Vulpes praecorsac (corrected herein) Ursidae Ursus deningeri (added herein) Ursus deningeri (den site) Euarctos thibetanus mediterraneus Ursus spelaeus “spelaeus” Mustelidae Martes sp. Mustela sp. Cervidae Cervidae Alces latifrons (added herein) Megaloceros antecedens (added herein) ?Praemegaceros/Megaloceros Capreolus sp. Suidae Sus scrofa subsp. (corrected herein) Equiidae Equus mosbachensis Bovidae Bovidae Bos primigenius subsp. Bos/ Bison schoetensacki (added herein) Rhinoceratidae hemitoechus Stephanorhinus kirchbergensis Proboscidae Palaeoloxodon antiquus (added herein) Castoridae Castor fiber Trogontherium cuvieri

P4 tooth impact mark at the trochlea, which fits exactly to a H. h. mosbachensis bit into the lion humerus (Fig. 4A). The production of a very typical hyena-like diagonal cut, due to cutting the joint for dismembering the lower leg, can be compared to a Late Pleistocene (C. c. spelaea) cut on a steppe lion humerus (P. l. spelaea) (cf. Diedrich, 2011c). The same stripped hyena breaking scissor dentition fits then again (another possible candidate is the spotted hyena C. c. praespelaea/ultima) into the diagonal cut of the distal bovid tibia (Fig. 4B), which would be the second indirect prove for its short-time scavenging presence at the Schoningn€ site. In addition to the typical large hyenid bite marks on both, the bovid tibia and the felid humerus, the lion humerus also has very small 1e2 mm sized triangular bite impacts over larger parts of its shaft. Those were interpreted as anthropogenic “hit marks” as result of its use as a “Neanderthal bone tool” by Serangeli et al. (2015). In fact, these small triangular impacts are only marks of the molar teeth of medium-sized to smaller carnivores, especially their upper jaw M1 3 triangular coned teeth (Fig. 4C). The best candidates for these small bite impacts on the humerus shaft includes especially the Fig. 5. Middle Pleistocene Parahyaena brunnea mosbachensis chewing on € Middle Pleistocene Cuon alpinus spelaeus, which is well known by a lion carcass at the northern German Holsteinian Interglacial (MIS 9e) Schoningen Lake shore site. In the background the saber-tooth cat Homotherium latidens older cub bone material from the nearby Holsteinian to Saalian Complex © (with cub fur ornamentation) trying to compete (artwork R. Uchytel). dated Baumann's Cave cave bear and Cuon den site (Diedrich, coll. Menger) which Upper jaw M1/P4 fits exactly in outline, morphology and size into the tooth bite marks of the of the trochlea. This makes sense, because hyenas use the breaking scissor teeth (see bite mark types and dentition in B) to dismember the leg parts and to cut of the joints. The Middle Pleistocene hyena most probably was the initial scavenger of a lion carcass which cracked the bone and chew damaged it on the distal lateral joint. The small triangular impact marks of the shaft “believed to be hit marks” are indeed well to identify as tooth impact marks of the red wolf Cuon alpinus subsp. (or small fox Vulpes praecorsac), which is found even in the Middle Pleistocene cave fauna of the Baumann's Cave 30 km far from the Schoningen€ archaeological Neanderthal camp/hunting site. B. Distal tibia of a bovid from the Middle Pleistocene (MIS 9e) Schoningen€ lake site (from Berkholst, 2015), which shows B. The same diagonal cut and even better as the lion humerus (cf. Fig. 3A) more complete breaking scissor dentition impact tooth marks (outlines and tooth tips) of a hyena herein used the C. Dentition and bite marks of brown extinct hyena P. b. mosbachensis. 82 C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83

2013a). Another and smaller possible candidate is the Middle Pleistocene Homotherium latidens with new records recorded Pleistocene smaller fox (Vulpes praecorsac), which is listed in the herein: Balve Cave in the Sauerland Karst, Zoolithen Cave in the Schoningen€ fauna as “small red fox Vulpes sp.” (cf. Kolfschoten, Frankonian Karst, Ketsch Rhine River Terrace Upper Rhine Graben. 2014). The new Mid-Pleistocene H. latidens humeri are discussed along In any case, there is no reason to believe that the “Homotherium” with the northern German Harz foreland Schoningen€ lake site humerus is from a saber-tooth cat, nor are its triangular impact incomplete felid humerus, which is revised to a smaller-sized lion marks caused due to “tool use” by humans. This lion humerus is or female due to absence of the supracondylar ridge. The German simply the product of interaction between carnivores (Fig. 5). A saber-tooth cat H. latidens records seem to reflect “dispersal events” possible scenario is illustrated herein: a lion carcass e perhaps within interglacials, several times in the Cromerian Complex MIS killed by hyenas (it can not be excluded to have been killed by during the MIS 19 Interglacial (Mosbach, West Runton), the MIS 15 Neanderthals with spear weapons) e as a result of competing with and 11 as shown for Czech cave (Sluj IV) records, and in the MIS 9e humans over the remains of hunted horses, which was finally Holsteinian Interglacial, (Schoningen).€ Probably, saber-tooth cats scavenged by brown hyenas, which might have killed a Homo- appeared again in the early Late Saalien two warm periods, the therium older cub at this site that tried to compete about the carcass Wackenian and Lietian Interstadials (MIS 7a, c), but they reached (Fig. 5). not into the final Saalian (MIS 6) nor survived into the Late Pleis- tocene in Europe. No saber-tooth cat is proven from an archaeo- 3.4. The big predators in competition e lions--- logical “killing” context. All known saber-tooth cat remains are hyenas-red found in hyena den cave/open air sites or in carnivore conflict contexts, including the Schoningen€ Lake site. Bite marks on a bovid The carnivores of the Middle Pleistocene in Germany and tibia and lion humerus from the Schoningen€ site can be demon- Europe are in need of a larger-scaled taxonomic, taphonomic and strated to have the same hyena breaking scissor tooth impact marks ethologic revision, but at least five larger felids were in competition on their distal joints. Hyenas produced similar “diagonal cuts” at like Panthera leo fossilis, Panthera onca gombaszoegensis, Panthera any guilt between joints for dismembering large body pieces (e.g. pardus sickenbergi, Acinonyx pardiensis intermedius, and Homo- legs) to move those from the site to avoid further carnivore or even therium latidens (cf. Hemmer, 2004; Hemmer et al., 2011; Diedrich, human conflicts. Those bone damages were left most probably at 2013c; Cherin et al., 2014; Anton et al., 2014). Lions were present in Schoningen€ and other Mid-Pleistocene sites by the early brown the late Middle Pleistocene MIS 9e as demonstrated herein by the hyena Parahyaena brunnea mosbachensis (or spotted hyena Crocuta small-sized lion humerus from Schoningen,€ which can be attrib- crocuta praespelaea/ultima), which is indirectly proven by its bite uted only to Panthera leo subsp. at the moment (most probably marks, and coprolites, and possibly directly by a scapula. female), but not to the much larger P. l. fossilis (cf. Dietrich, 1968). This humerus is also far too large for P. o. gombaszoegensis. We can References only speculate on the presence of this (e.g. Hemmer et al., 2003) and the cheetah Acinonyx pardinensis intermedius (e.g. Adam, K.D., 1975. Die mittelpleistozane€ Saugetier-Fauna€ aus dem Heppenloch bei € Hemmer et al., 2011) at this time in the Schoningen/Harz€ Mountain Gutenberg (Württemberg). Stuttg. Beitr. Naturkd. Ser. B Geol. Palaontol. 3, 1e247. foreland region. Similar in size as modern forms were the early Anton, M., Galobart, A., Turner, A., 2005. Co-existence of scimitar-toothed cats, lions spotted hyenas described from a skull and scant material from the and hominins in the European Pleistocene. Implications of the post-cranial early Middle Pleistocene Mosbach site (Reichenau, 1906; Schütt, anatomy of Homotherium latidens (Owen) for comparative palaeoecology. fi Quat. Sci. Rev. 24 (10/11), 1287e1301. 1971). A new nd of an incomplete skull from the Middle Pleisto- Anton, M., Salesa, M., Turner, A., Galobart, A., Pastor, J., 2009. Soft tissue recon- cene of the Rhine Graben (Fig. 4C) has similar dentition to the early struction of Homotherium latidens (Mammalia, , Felidae). Implications brown hyena Parahyaena brunnea mosbachensis (Schütt, 1971). for the possibility of representations in Palaeolithic art. Geobios 42 (5), e Possibly, an undescribed scapula from Schoningen€ of an older in- 541 551. Anton, M., Salesa, M.J., Galobart, A., Tseng, Z.J., 2014. The Plio-Pleistocene scimitar- dividual (with pathology) belongs to H. h. mosbachensis, which toothed felid genus Homotherium Fabrini, 1890 (, Homo- improves directly the carnivore biodiversity and presence of those therini): diversity, palaeogeography and taxonomic implications. Quat. Sci. Rev. striped hyenas at Schoningen€ (Table 1). Another hyena of this time 96, 259. Cherin 268. Ballesio, R., 1963. Monographie d’un du gisement villafranchien de is an early spotted hyena Crocuta crocuta praespelaea (Schütt, 1971) Seneze: Homotherium crenatidens Fabrini. Trav. Lab. Geol. Lyon Nouv. Ser. 9, which was also found in the Mosbach carnivore fauna. Those two 1e129. Berkholst, B.E., 2015. The large fauna of the Pleistocene site Schoningen€ hyenids from the Middle Pleistocene of Europe must have € € 13II. The levels Scho 13II-1, 13II-2 and 13II-3 (Unpublished Master-Thesis). competed with all felids, especially Homotherium. At Schoningen, University Leiden. the presence of is indirectly proven by their bite Cherin, M., Iurino, D.A., Sardella, R., Rook, L., 2014. Acinonyx pardiensis (Carnivora, marks on prey bones, which are also found at modern stripped Felidae) from the Early Pleistocene of Pantalla (Italy): predatory behaviour and ecological role of the giant Plio-Pleistocene cheetah. Quat. Sci. Rev. 87, 82e97. hyena den site bone accumulations (cf. Owens and Owens, 1979; Dawkins, W.B., Sandford, W.A., 1872. The British pleistocene Mammalia. Part IV. Skinner and Van Aarde, 1991; Fourvel et al., 2015). An open air British Pleistocene Felidae: Felis pardus, Lin.; Felis caffer, Desm.; Felis catus, Lin.; hyena den site (striped or spotted hyena) along the Mid-Pleistocene Machaerodus latidens, Owen. € Diedrich, C., 2005. Von eiszeitlichen Fleckenhyanen€ eingeschleppte Reste des Schoningen Lake is not yet known, but at least some coprolites have Steppenwisents Bison priscus Bojanus, 1827 aus dem oberpleistozanen€ Fleck- been found (pers. com. Serangeli), which are the third indicator for enhyanenhorst€ des Perick-Hohlensystems€ (NW Deutschland). Philippia 12 (1), short-time presence. Spotted hyenas C. c. praespelaea/ultima seem 21e30. to have occupied during the late Mid-Pleistocene (Holsteinian/ Diedrich, C., 2009. Late Pleistocene (Acanthion) brachyura Linnaeus, 1758 from the Fuchsluken cave at the Rote Berg near Saalfeld (Thuringia, Germany) - Saalian) a former entrance at the only 30 km nearby Baumann's a porcupine and hyena den and contribution to their palaeobiogeography. Open Cave (Diedrich, 2013a). Palaeontol. J. 2008 (1), 33e41. Diedrich, C., 2011a. The largest European lion Panthera leo spelaea (Goldfuss) pop- ulation from the Zoolithen Cave, Germany e specialized cave bear predators of 4. Conclusions Europe. Hist. Biol. 23 (2e3), 271e311. Diedrich, C., 2011b. Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from In Germany, there are few sites with isolated tooth and post- the Balve Cave (NW Germany) e a cave bear, hyena den and Middle Palaeolithic human cave, and review of the Sauerland Karst lion sites. Quaternaire 22 (2), cranial bone remains of the Early Pleistocene Homotherium cren- 105e127. atidens saber-tooth cat (e.g. Untermassfeld), and the Mid- Diedrich, C., 2011c. Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 83

footprints and bone remains from open air sites in northern Germany e evi- Koutsodendris, A., Müller, U.C., Pross, J., Brauer, A., Kotthoff, U., Lotter, A.F., 2010. dence of hyena-lion antagonism in Europe. Quat. Sci. Rev. 30, 1883e1906. Vegetation dynamics and climate variability during the Holsteinian interglacial Diedrich, C., 2011d. Periodical use of the Balve Cave (NW Germany) as a Late based on a pollen record from Dethlingen (northern Germany). Quat. Sci. Rev. Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) den: Hyena occupations 29, 3298e3307. and bone accumulations vs. human Middle Palaeolithic activity. Quat. Int. 233, Lewis, M., Pacher, M., Turner, A., 2010. Carnivora from the west runton freshwater 171e184. bed. Quat. Int. 228, 116e135. Diedrich, C., 2012. The Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) McFarlane, D.A., Lundberg, J., 2013. On the occurrence of the scimitar-toothed cat, population from the Emscher River terrace hyena open air den Bottrop and Homotherium latidens (Carnivora; Felidae), at Kents Cavern, England. other sites in NW-Germany e woolly scavengers and their bone J. Archaeol. Sci. 40, 1629e1635. accumulations along rivers in lowland steppe environments. Quat. Müller, H., 1974. Pollenanalytische Untersuchungen und Jahresschichtenzahlungen€ Int. 276e277, 93e119. an der Holstein-zeitlichen Kieselgur von Munstere Breloh. Geol. Jahrb. A 21, Diedrich, C., 2013a. Evolution, Horste, Taphonomie und Pradatoren€ der Rübelander€ 107e140. Hohlenb€ aren,€ Harz (Norddeutschland). Mittl. Verb. Dtsch. Hohlen€ Karstforscher Nitychoruk, J., Binka, K., Hoefs, J., Ruppert, H., Schneider, J., 2005. Climate recon- 59 (1), 4e29. struction for the Holsteinian interglacial in eastern Poland and its comparison Diedrich, C., 2013b. Late Pleistocene hyena and steppe lion feeding strate- with isotopic data from marine isotope stage 11. Quat. Sci. Rev. 24, 631e644. gies on their largest preydPalaeoloxodon antiquus Falconer and Cautley 1845 at Owen, R., 1843. Report on the British Mammalia, Part 1. Brit. Assoc. Report for 1842, the straight-tusked elephant graveyard and Neanderthal site Neumark-Nord p. 68. Lake 1, Central Germany. Archaeol. Anthropol. Sci. 6 (3), 271e291. Owens, D.D., Owens, M.J., 1979. Communal denning and clan association in brown Diedrich, C., 2013c. Late Pleistocene across Europe - most northern Eu- (Hyaena brunnea Thunberg) of the central Kalahari Desert. Afr. J. Ecol. ropean population, highest elevated records in the Alps, complete skeletons in 17, 35e44. the Dinarids and comparison to the Ice Age cave art. Quat. Sci. Rev. 76, 167e193. Palmqvist, P., Martínez-Navarro, B., Perez-Claros, J.A., Torregrosa, V., Figueirido, B., Diedrich, C., 2014. Holotype skulls, stratigraphy, bone taphonomy and excavation Jimenez-Arenas, J.M., Espigares, M.P., Ros-Montoya, S., De Renzi, M., 2011. The history in the Zoolithen Cave and new theory about Esper's “great deluge”. giant hyena Pachycrocuta brevirostris: Modelling the bone-cracking behavior of Quat. Sci. J. 63 (1), 78e98. an extinct carnivore. Quat. Int. 243 (1), 61e79. Diedrich, C., Rathgeber, T., 2012. Late Pleistocene steppe lion Panthera leo spelaea Reichenau, W. v, 1906. Über einen Schadel€ der Hyaena avernensis Croizet et Jobert (Goldfuss, 1810) skeleton remains of the Upper Rhine valley (SW Germany) and aus dem Mosbacher Sande. Jahrb. Des. Nassau. Ver. Naturkd. 58, 175e182. contribution to their sexual dimorphism, taphonomy and habitus. Hist. Biol. 24 Reumer, J.W.F., Rook, L., Van Der Borg, K., Post, K., Mol, D., De Vos, J., 2003. Late (1), 1e28. Pleistocene survival of the saber-toothed cat Homotherium in northwestern Dietrich, W.O., 1968. Fossile Lowen€ im europaischen€ und afrikanischen Pleistozan.€ Europe. J. Vertebr. Paleontol. 23, 260e262. Palaontol.€ Abh. A Palaozool.€ 3 (2), 323e366. Sardella, S., Iurino, D.A., 2012. The latest Early Pleistocene sabertoothed cat Fejfar, O., 1956. Seznam druhu fosilních savcu z jeskyne C 718 na Zlatem koni u Homotherium (Felidae, Mammalia) from Monte Peglia (Umbria, central Italy). Koneprus. Vestník Ústred. ústavu Geol. Praze 31, 274e276. Boll. Soc. Paleontol. Ital. 51 (1), 15e22. Fourvel, J.B., Avery, G., Fosse, P., 2015. Spotted, striped or brown? Taphonomic Schütt, G., 1970. Nachweis der Sabelzahnkatze€ Homotherium in den Altpleistozanen€ studies at dens of extant hyaenas in eastern and southern . Quat. Int. 369, Mosbacher Sanden (Wiesbaden, Hessen). Neues Jahrb. Geol. Palaontol.€ Mon- 38e50. atsh. 3, 187e192. Frewein, J., Vollmerhaus, B., 1994. Anatomie von Hund und Katze. Blackwell Wis- Schütt, G., 1971. Die Hyanen€ der Mosbacher Sande (Altpleistozan,€ Wiesbaden/ senschaftsverlag, p. 457. Hessen) mit einem Beitrag zur Stammesgeschichte der Gattung Crocuta. Mainz. Galobart, A., Pons-Moya, J., Anton, M., Maroto, J., 2003. Descripcion del material de Naturwiss. Arch. 10, 29e76. Homotherium latidens (Owen) de los yacimentos del Pleistoceno inferior de Serangeli, J., Kolfschoten, T., von Starkovich, B.M., Verheijen, I., 2015. The European Incarcal (Girona, NE de la Penísula Iberica). Paleontol. I Evol. 34, 99e141. saber-toothed cat (Homotherium latidens) found in the “Spear Horizon” at Hemmer, H., 2004. About scimitar cats and cave lions e rise and fall of Pleistocene Schoningen€ (Germany). J. Hum. Evol. 89, 172e180. life-communities over the continents. In: Abstract 18th Internnational Senck- Skinner, J.D., Van Aarde, R.J., 1991. Bone collecting by brown hyaenas Hyaena enberg Conference, Weimar, pp. 1e2. brunnea in the central Namib Desert, Namibia. J. Archaeol. Sci. 18, 513e523. Hemmer, H., Kahlke, R.-D., Keller, T., 2003. Panthera onca gombaszoegensis (Kretzoi, Stephan, H.-J., 2014. Climato-stratigraphic subdivision of the Pleistocene in 1938) aus den frühmittelpleistozanen€ Mosbach-Sanden (Wiesbaden, Hessen, Schleswig-Holstein, Germany and adjoining areas e status and problems. Eis- Deutschland) e ein Beitrag zur Kenntnis der Variabilitat€ und Ver- zeitalter und Gegenwart. Quat. Sci. J. 63 (1), 3e18. breitungsgeschichte des Jaguars. Neues Jahrb. Geol. Palaontol.€ 229 (1), 31e60. Stuart, A.J., Lister, A., 2010. The west runton freshwater bed and the west runton Hemmer, H., Kahlke, R.D., Vekua, A.K., 2011. The cheetah Acinonyx pardinensis mammoth: summary and conclusions. Quat. Int. 228 (1), 241e248. (Croizet et Jobert, 1828) s.l. at the hominin site of (Georgia) - a po- Tseng, Z.J., Chang, C.-H., 2007. A Study of New Material of Crocuta crocuta ultima tential prime meat supplier in Early Pleistocene ecosystems. Quat. Sci. Rev. 30 (Carnivora: Hyaenidae) from the Quaternary of Taiwan. Collect. Res. 20, 9e19. (19e20), 2703e2714. Weithofer, K.A., 1889. Die fossilen Hyaenen des Arnothals. Denkschr. Akad. Wiss. Kolfschoten, T. van, 2014. The Palaeolithic locality Schoningen€ (Germany): a review Math. Naturwissenschaftliche Kl. Wien 55. of the mammalian record. Quat. Int. 326/327, 469e480.