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Quaternary International 436 (2017) 76e83

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Quaternary International

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Homotherium from Middle Pleistocene archaeological and carnivoreden sites of Germany e Taxonomy, taphonomy and a revision of theSch€oningen, West Runton and other saber-tooth cat sites

Cajus G. Diedrich a, b, *, 1, Donald A. McFarlane c

a PaleoLogic, Private Research Institute, Petra Bezruce 96, CZ-26751 Zdice, Czech Republicb University of Koblenz-Landau, IfIN, Department of Biology, Universit€atsstr. 1, D-56070 Koblenz, Germanyc Keck Science Department, The Claremont Colleges, 925 North Mills Avenue, Claremont, CA 91711-5916, USA

a r t i c l e i n f o

Article history:Received 19 April 2016Received in revised form7 October 2016Accepted 13 October 2016Available online 1 March 2017

Keywords:HomotheriumHolsteinian Interglacial (MIS 9e)GermanyArchaeological camp versus hyena dencontextHyaena and Cuon bite damageLion versus saber tooth cat humeriNo human tool

* Corresponding author. PaleoLogic, Private ResearcCZ-26751 Zdice, Czech Republic.

E-mail addresses: cdiedri@gmx.net (C.G. Diedclaremont.edu (D.A. McFarlane).

1 www.paleologic.eu.

http://dx.doi.org/10.1016/j.quaint.2016.10.0151040-6182/© 2016 Elsevier Ltd and INQUA. All rights

a b s t r a c t

Four new saber-tooth cat (Homotherium) sites in Germany with new dental and postcranial bone ma-terial are different in their taphonomic context: 1. The Archaeological Middle Palaeolithic (MIS 9e-Interglacial) Sch€oningen Lake site with remains of a cub carcass, 2. The Middle Palaeolithic (MIS 5e-9)Archaeological/cave bear den site of Balve Cave yielding a lower canine tooth of an older individual, 3.The Zoolithen Cave (MIS 3e9) cave bear/hyena den with one distal half humerus of an adult, 4. TheKetsch open air Rhine River terrace site which has provided another distal humerus of an adult saber-tooth cat. Whereas only the Sch€oningen site is precisely dated as Holsteinian Interglacial (approx.330.000e315.000 BP), all other material seems to come from the same Middle Pleistocene warm period,or few younger Saalian interstadials (MIS 7a, e) deposits, and did not extend over the last MIS 7 glacialinto the Late Pleistocene. Homotherium as hyena-like slow moving cat seems to have disappeared withinthe Saalian due to competition with other scavengers like Ice Age spotted and brown hyenas (Crocutacrocuta praespelaea/ultima and Pachycrocuta brunnea mosbachensis). The juvenile saber-tooth cat cubfrom Sch€oningen might be in archaeological context or represent only a carnivore kill. At the ZoolithenCave, the single bone must have been imported into a hyena prey bone assemblage. The situation ispossibly similar at the two other sites Ketsch and Balve Cave. The formerly described Sch€oningen “saber-tooth cat” humerus is revised, such as other opposite as lion humeri described material from differentEuropean sites. The presence of the well-developed supracondylar ridge distinguishes Homotheriumwellfrom Middle/Late Pleistocene lions Panthera leo (e.g. spelaea, fossilis). The Sch€oningen lion humrus hasbeen chew-cut first most probably by a stripped hyena whose cutting scissor teeth produced a diagonalbite cut and P4/M1 impact marks around the trochlea. 1e2 mm small, mostly triangular-oval bite markson the lion humerus shaft compacta results from a second scavenger and not from “Neanderthal tooluse”. Those bite mark sizes are produced mainly of the upper molar teeth of a the red wolf Cuon alpinussubsp. (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

The only known European saber-tooth cat Homotherium cren-atidens (Weithofer, 1889) skeleton from the Early Pleistocene (Vil-lafranchian) of the French site S�eneze is well published in a

h Institute, Petra Bezruce 96,

rich), DMcFarlane@kecksci.

reserved.

monograph by Ballesio (1963). This specimen includes completepreserved humeri, which are compared herein for a revision ofseveral questionable “distal humeri” described in recent years fromseveral Central European sites often as “felid or lion humeri” (e.g.Lewis et al., 2010). The Middle Pleistocene European saber-toothcats, which are not yet known by skeletons, are assigned toHomotherium latidens (Owen, 1843) (e.g. Ant�on et al., 2005), forwhich one of the most northern records comes from an Englishcave (Dawkins and Sandford, 1872; McFarlane and Lundberg, 2013).The “Late Pleistocene” records from the North Sea (Reumer et al.,2003) are without stratigraphic value or reliable dates because of

C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 77

their embedding in modern marine sediments: no other Europeanfinds support their survival into the last Ice Age.

Thematerial found in Germany (Fig.1A) is quite sparse and lackseven complete skulls. A few single teeth and postcranial elementshave been published. The Early Pleistocene records are from theriver terrace site Untermabfeld (cf. Hemmer et al., 2011). Somematerial comes from the Mosbach Sands Neckar River terrace(Schütt, 1970), which are revised by the actual stratigraphy of thissite (cf. Stephan, 2014) now into the early Middle PleistoceneCromerian Complex. Further Middle Pleistocene material wasfound in the “Heppenloch” also being called Gutenberg Cave(Adam, 1975).

In this contribution, Middle Pleistocenematerial from three newGerman sites can be added (Figs. 1e3). It is not intended to be a fullreview of German, or revision of Eurasian Pleistocene saber-toothcat sites, but it is of urgent need because material has beenconfused. Some saber-tooth cat remains are figured (only 0,1%, ineach publication) in lion publications (cf. Diedrich, 2011a,b;Diedrich and Rathgeber, 2012). Opposite, some lion remains havebeen misidentified as saber-tooth cat remains. For example, thenewest “saber-tooth cat” records described by Serangeli et al.(2015) represent only one saber-tooth cat cub dentition and to itsage class fitting humerus such as few other bone fragments of a cub(not two individuals as discussed therein). We revise the figuredhumerus determined in their study as “Homotherium” (second in-dividual) here. Furthermore, the well-known taphonomy of Euro-pean Late Pleistocene lion carcasses caused mainly by hyenafeeding (Diedrich, 2011a) can be extended with more details of

Fig. 1. Pleistocene saber-tooth cat Homotherium sites in Germany and Czech Republic and ot1975; Stuart and Lister, 2010; Serangeli et al., 2015; with new German sites being added).

hyena feeding strategies on their felid competitors, lions and saber-tooth cats.

2. Material and methods

The carnivore material of the Paleon - Sch€oningen SpearMuseum (¼ Pal) was only screened and is partly exposed in thepermanent exhibition, which needs further detailed work in future.The humerus discussed herein was checked by the tooth markforms and sizes. The studied saber-tooth cat and other bone ma-terial of the Balve Cave is stored in the Geologisch-Pal€aontologischeMuseum of the Westphalian Wilhelms-University Münster (¼GPIM). The Zoolithen Cave material is in the collection of theFr€ankische H€ohlen und Karstforscher Club (¼ FHKF). One humerusfragment is stored in the Staatliche Museum für NaturkundeStuttgart (¼ SMNS). One complete lion humerus is stored in theLandesamt Sachsen Anhalt collection (¼ LDA). The upper jaw saber-tooth cat canine from the Czech C718 Cave site is in the Natio-nalmuseum Prague (¼ NMP). The compared damaged lion hu-merus from the Neander Valley is in the Goldfuss-Museum of theRheinischeWilhelms-University Bonn (¼ GMB). A single tooth cuspfrom Sanderhausen is kept in the collection of the Spenglermu-seum Sangerhausen (¼ SPM).

2.1. Geology and dating

All new German sites are of Middle Pleistocene (Cromerian-Saalian, ¼ MIS 21-7) in age according to the updated German

her compared sites in England (dating composed from Fejfar, 1956; Schütt, 1970; Adam,

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 catfrom the Konĕprusy Doline, Bohemian Karst, Czech Republic (coll. NMP). 2. H. latidens, left lower jaw canine of an older cat from the Balve Cave, Sauerland Karst, Germany (modifiedfrom 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 serratedcutting edges and non-closed roots) of an older cub (half to maximum one year) from Sch€oningen Lake shore site, Germany (modified from Serangeli et al., 2015): 5. Caninefragment. Right I3, 6. Left P4, 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.

C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e8378

stratigraphic chart (cf. Stephan, 2014; cf. Fig. 1B). The H. latidens re-mains from Sch€oningen are dated according to the newest work onthe Middle to Late Pleistocene stratigraphy in northern Germany, in

Fig. 3. Early-Middle Pleistocene saber-tooth cat Homotherium humeri from European sites.(from Ballesio, 1963), cranial. 2. Humerus from Incarcal (Es) (from Ant�on et al., 2005), cranial.West Runton (GB) (from Lewis et al., 2010). 5. Left distal half humerus of a grown up from thehalf humerus of a grown up cat from the Rhine River Terrace Ketsch-Kreuzwiese (coll. SMN

which the Holsteinian Interglacial is dated into the MIS 9e isotopestage (330.000e315.00 BP) (Müller, 1974). It is characterized by thedevelopment in general of temperate mixed forests in central and

1. Humerus (mirrored) from the H. crenatidens (Weithofer, 1889) skeleton of Sen�eze (F)3. Distal humerus from C718 Cave (CZ) (coll. NMP), caudal. 4. Right distal humerus fromZoolithen Cave (D), cranial and mirrored (modified from Diedrich, 2011d). 6. Left distalS), cranial.

C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 79

northwestern Europe and seasonality (e.g., Nitychoruk et al., 2005;Koutsodendris et al., 2010). The Zoolithen Cave has been recentlyrevised and its bone remains are thought to have been accumulatednot only in Late Pleistocene times, but instead since the Elsterian/Holsteinian (Diedrich, 2014). The long-termmultiple use of this caveas a cave bear, hyena and wolf den within different climatic periodsexplains the large amount of bones (mainly different cave bear spe-cies/subspecies of Middle Pleistocene: Ursus deningeri, Late Pleisto-cene: Ursus spelaeus eremus/spelaeus, Ursus ingressus) and thepresence of the “largest amount of lion remains ever excavated in aEuropean cave”, which also includes Middle Pleistocene P. l. fossilisremains (Diedrich, 2011a, 2014). All material, including the raresaber-tooth remains, are in secondary/tertiary positions within thecave system due to predator activities and later floods and can not bedated exactly in many cases, except using biostratigraphy (e.g.C. alpinus spelaeus and H. latidens are such as U. deningeri MiddlePleistocene species: Diedrich, 2014). The third site discussed here,Ketsch, is a Rhine River Valley gravel pit terrace site, which hasmixedMiddle tomainly Late Pleistocenemegafauna elements (Diedrich andRathgeber, 2012).

3. Results and discussion

3.1. Saber-tooth cat Homotherium versus lion Panthera humerusosteology

In most previous accounts of Middle Pleistocene H. latidens, thedistal humeri were figured, but in no case was the most importantdefining character discussed, although it was also used in soft tissuefore limb reconstructions by Ant�on et al. (2009). Based on the com-plete and well-preserved Plio-/Early Pleistocene H. crenatidens hu-merus figured by Ballesio (1963) (Fig. 3.1), the supracondylar ridge isabsent in this genus, which does not allow a rotation of the paw (cf.Ant�on et al., 2009). Here, in one example figured (Fig. 4A), for a smalllion (most probaly female) from the Late Pleistocene of the Her-mann's Cave (Harz Mountain Range), the ridge is strong and about2e3 mm wide e a level of development similar to that seen inmodern and Middle to Late Pleistocene lions (P. l. leo, P. l. spelaea, P. l.fossilis: cf. Dietrich,1968; Diedrich, 2011a,b). On this ridge, the rotatormusculature (Brachiradialis) for the lower forelimb is attached to theradius (e.g. Frewein and Vollmerhaus, 1994; Ant�on et al., 2009)(Fig. 4A). This allows lions to catch its prey with the forelimb paws(Ant�on et al., 2009). It is much different in the saber-tooth catHomotherium (as in hyenids, canids), where absence of the supra-condylar ridge does not allow the forelimbs to be rotated and used inprey capture (Ant�on et al., 2009). Also, based on this osteologicalcharacter, it is likely that the gait of Homotheriumwas “hyenae like”(Ant�on et al., 2005, 2009, 2014). The supracondylar ridge is also ab-sent in the two distal humeri from the Zoolithen Cave and Ketschriver terrace site (Fig. 3). Using this very important ostelologic char-acter, the felid humerus from Sch€oningen figured by Serangeli et al.(2015) (Fig. 4A) is a typical lion humerus by comparison to theH. crenatidens holotype humerus (cf. Ballesio, 1963: Fig. 3). Interest-ingly, the length of the supracondylar ridge is 5e10mmshorter in theSch€oningen specimen than in other Middle-Late Pleistocene lions, issome such as the Late Pleistocene Hermann's Cave lion humerus(Fig. 4A). The Mid-/Late Pleistocene lions from Germany are still inrevision about this theme of “three different sized” forms (not vari-ability nor sexual dimorphism) are not discussed herein further.

3.2. The humeri osteologic and biostratigraphic differences inH. crenatidens/latidens

The material from Germany is few and most is incomplete, butthe humeri allow a new discussion to the validities of the Early

H. crenatidens andMiddle PleistoceneH. latidens species (Figs.1 and3). The humerus differences of both biostratigraphic relevant spe-cies seem not to relate to sexual dimorphism or variability in sizes,as thought to have been demonstrated for Pliocene saber-tooth catmandibles of Europe and Northern America (Ant�on et al., 2014). Asa simple fact of a strong faunal turnover between the Early/Mid-Pleistocene, it must be expected, that there was also a specieschange/evolutionary trend within Homotherium, similar as forother felids (e.g. Hemmer, 2004).

The Early Pleistocene holotype skeleton of H. crenatidens isrecently under new studies, and original material was not possibleinclude. The most obvious character to distinguish the humeri ofH. crenatidens and H. latidens lies in the difference of the complete,and especially the distal shaft width, which is about 5 mm largerand more robust in the older H. crenatidens (Fig. 3). Based on theEarly Pleistocene Sen�eze skeleton H. crenatidens humerus, the EarlyPleistocene Homotherium humerus figured from the Spanish siteIncarcal described as “H. latidens” (cf. Galobart et al., 2003; Ant�onet al., 2005) must be revised herein to belong to H. crenatidens,also correlating then with the biostratigraphy (Fig. 1B). Also Italianfinds (cf. Sardella and Iurino, 2012) seem to belong to this olderspecies.

The humeri from the Middle Pleistocene H. latidens are slimmeror gracile in their shafts in all herein figured English and GermanMiddle Pleistocene specimens (Fig. 2). The comparison between thehumeri figured herein (Fig. 3) includes the “P. leo” humerus fromthe Cromerian West Runton interglacial MIS 19 (Stuart and Lister,2010) site in England (cf. Lewis et al., 2010). This humerus is,compared to the H. crenatidens holotype humerus in same size andshape in the distal shaft width to the German finds (Fig. 2). TheEnglish West Runton site also delivered a calcaneus of H. latidens(Lewis et al., 2010), which finally supports the humerusidentification.

3.3. Hyena bone damage versus human “bone tools”

The taphonomy of carnivore bone damage for the Sch€oningensite is misinterpreted in both papers of Berkholst (2015) andSerangeli et al. (2015). Furthermore, the carnivores were not yetwell determined (c.f Kolfschoten, 2014), but are well known now inmost cases (see Table 1). Older literature and important materialhas been overlooked, for example the only complete stripped hyenaancestor skull of Parahyaena brunnea mosbachensis (cf. Reichenau,1906; Schütt, 1971), to which material another partial skull canbe added from the Middle Pleistocene of the Upper Rhine Grabenherein as very rare record in Central Europe (Fig. 3). The spottedand brown hyenas are still to be discussed, if those are late EarlyPleistocene Parahyaena brunnea mosbachensis (in some casesPachycrocuta: e.g. Palmqvist et al., 2011) or Crocuta crocuta praes-pelaea/ultima (cf. Schütt, 1971; Tseng and Chang, 2007) forms in theMid-Pleistocene. Whereas Berkholst (2015) identified bite damageon a distal bovid tibia as “rodent bite marks”, those should havebeen compared to the larger rodent bite marks of species such asPleistocene porcupines, which are completely different (cf.Diedrich, 2009). The bite marks on this bovid guilt bone (seeFig. 4B) are in fact typical large predator bite marks, based on bitemark analysis of large prey (e.g elephants, rhinoceroses and evenbisons: Diedrich, 2005, 2012, 2013b). Instead of classical “bite markmetric” analyses, the much more successful direct digital-projection test of the “bite-fit” into the bite marks of differentpredator candidates was applied to the lion humerus and the bovidtibia from Sch€oningen (Fig. 4B). The test of the largest predatorsHomotherium, Panthera, Hyeana and Crocuta dentitions demon-strates that in both cases only hyenas account for all the bitedamage (see Fig. 4). Most convincing is the fit of the cutting scissor

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 Sch€oningen lake in comparison to similar sized lion humeri from LatePleistocene German caves (1. Hermann's Cave cave bear den, 2. Teufelskammer Cave hyena den). The typical lion character (Supracondylar ridge) and also much thicker and not ovalshaft (cf. differences in Homotherium humeri Fig. 3) let identify this humerus fragment as a lion subspecies. Furthermore, the taphonomic felid humeri comparison demonstrate thechew/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) 76e8380

Table 1Updated megafauna list of the Middle Pleistocene Sch€oningen Pre-Neanderthal hunter camp and Homotherium lake shore site with “mixedforest/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).

Sch€oningen Lake shoreHarz Foreland (MIS 9e Interglacial)

Baumann's CaveHarz Mountain Range (MIS 6e9 Stadials/Interstadials)

Felidae FelidaePanthera leo subsp. (small lion)(added herein)

Panthera leo subsp. (small lion)

Homotherium latidens (corrected herein) Panthera leo fossilis (large lion)Hyaenidae HyaenidaeParahyeana brunnea cf. mosbachensis sp. (added herein) Crocuta crocuta praespelaea/ultimaCanidae CanidaeCuon alpinus spelaeus (added herein) Cuon alpinus spelaeusVulpes praecorsac (corrected herein)UrsidaeUrsus deningeri (added herein) Ursus deningeri (den site)Euarctos thibetanus mediterraneus Ursus spelaeus “spelaeus”MustelidaeMartes sp.Mustela sp.Cervidae CervidaeAlces latifrons (added herein)Megaloceros antecedens (added herein) ?Praemegaceros/MegalocerosCapreolus sp.SuidaeSus scrofa subsp. (corrected herein)EquiidaeEquus mosbachensisBovidae BovidaeBos primigenius subsp. Bos/BisonBison schoetensacki (added herein)RhinoceratidaeStephanorhinus hemitoechusStephanorhinus kirchbergensisProboscidaePalaeoloxodon antiquus (added herein)CastoridaeCastor fiberTrogontherium cuvieri

Fig. 5. Middle Pleistocene brown hyena Parahyaena brunnea mosbachensis chewing ona lion carcass at the northern German Holsteinian Interglacial (MIS 9e) Sch€oningenLake shore site. In the background the saber-tooth cat Homotherium latidens older cub(with cub fur ornamentation) trying to compete (artwork © R. Uchytel).

C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e83 81

P4 tooth impact mark at the trochlea, which fits exactly to a H. h.mosbachensis bit into the lion humerus (Fig. 4A). The production ofa very typical hyena-like diagonal cut, due to cutting the joint fordismembering the lower leg, can be compared to a Late Pleistocenespotted hyena (C. c. spelaea) cut on a steppe lion humerus (P. l.spelaea) (cf. Diedrich, 2011c). The same stripped hyena breakingscissor dentition fits then again (another possible candidate is thespotted hyena C. c. praespelaea/ultima) into the diagonal cut of thedistal bovid tibia (Fig. 4B), which would be the second indirectprove for its short-time scavenging presence at the Sch€oningn site.In addition to the typical large hyenid bite marks on both, the bovidtibia and the felid humerus, the lion humerus also has very small1e2 mm sized triangular bite impacts over larger parts of its shaft.Those were interpreted as anthropogenic “hit marks” as result of itsuse as a “Neanderthal bone tool” by Serangeli et al. (2015). In fact,these small triangular impacts are only marks of the molar teeth ofmedium-sized to smaller carnivores, especially their upper jawM1�3 triangular coned teeth (Fig. 4C). The best candidates for thesesmall bite impacts on the humerus shaft includes especially theMiddle Pleistocene Cuon alpinus spelaeus, which is well known bybone material from the nearby Holsteinian to Saalian Complexdated 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 breakingscissor 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 alion 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 toidentify 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 Cave30 km far from the Sch€oningen archaeological Neanderthal camp/hunting site. B. Distal tibia of a bovid from the Middle Pleistocene (MIS 9e) Sch€oningen 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.

C.G. Diedrich, D.A. McFarlane / Quaternary International 436 (2017) 76e8382

2013a). Another and smaller possible candidate is the MiddlePleistocene smaller fox (Vulpes praecorsac), which is listed in theSch€oningen fauna as “small red fox Vulpes sp.” (cf. Kolfschoten,2014).

In any case, there is no reason to believe that the “Homotherium”

humerus is from a saber-tooth cat, nor are its triangular impactmarks caused due to “tool use” by humans. This lion humerus issimply the product of interaction between carnivores (Fig. 5). Apossible scenario is illustrated herein: a lion carcass e perhapskilled by hyenas (it can not be excluded to have been killed byNeanderthals with spear weapons) e as a result of competing withhumans over the remains of hunted horses, which was finallyscavenged by brown hyenas, which might have killed a Homo-therium older cub at this site that tried to compete about the carcass(Fig. 5).

3.4. The big predators in competition e lions-jaguars-cheetah-hyenas-red wolves

The carnivores of the Middle Pleistocene in Germany andEurope are in need of a larger-scaled taxonomic, taphonomic andethologic revision, but at least five larger felids were in competitionlike Panthera leo fossilis, Panthera onca gombaszoegensis, Pantherapardus sickenbergi, Acinonyx pardiensis intermedius, and Homo-therium latidens (cf. Hemmer, 2004; Hemmer et al., 2011; Diedrich,2013c; Cherin et al., 2014; Ant�on et al., 2014). Lions were present inthe late Middle Pleistocene MIS 9e as demonstrated herein by thesmall-sized lion humerus from Sch€oningen, which can be attrib-uted only to Panthera leo subsp. at the moment (most probablyfemale), but not to the much larger P. l. fossilis (cf. Dietrich, 1968).This humerus is also far too large for P. o. gombaszoegensis. We canonly speculate on the presence of this jaguar (e.g. Hemmer et al.,2003) and the cheetah Acinonyx pardinensis intermedius (e.g.Hemmer et al., 2011) at this time in the Sch€oningen/Harz Mountainforeland region. Similar in size as modern forms were the earlyspotted hyenas described from a skull and scant material from theearly Middle Pleistocene Mosbach site (Reichenau, 1906; Schütt,1971). A new find of an incomplete skull from the Middle Pleisto-cene of the Rhine Graben (Fig. 4C) has similar dentition to the earlybrown hyena Parahyaena brunnea mosbachensis (Schütt, 1971).Possibly, an undescribed scapula from Sch€oningen of an older in-dividual (with pathology) belongs to H. h. mosbachensis, whichimproves directly the carnivore biodiversity and presence of thosestriped hyenas at Sch€oningen (Table 1). Another hyena of this timeis an early spotted hyena Crocuta crocuta praespelaea (Schütt, 1971)which was also found in the Mosbach carnivore fauna. Those twohyenids from the Middle Pleistocene of Europe must havecompeted with all felids, especially Homotherium. At Sch€oningen,the presence of striped hyena is indirectly proven by their bitemarks on prey bones, which are also found at modern strippedhyena den site bone accumulations (cf. Owens and Owens, 1979;Skinner and Van Aarde, 1991; Fourvel et al., 2015). An open airhyena den site (striped or spotted hyena) along theMid-PleistoceneSch€oningen Lake is not yet known, but at least some coprolites havebeen found (pers. com. Serangeli), which are the third indicator forshort-time presence. Spotted hyenas C. c. praespelaea/ultima seemto have occupied during the late Mid-Pleistocene (Holsteinian/Saalian) a former entrance at the only 30 km nearby Baumann'sCave (Diedrich, 2013a).

4. Conclusions

In Germany, there are few sites with isolated tooth and post-cranial bone remains of the Early Pleistocene Homotherium cren-atidens saber-tooth cat (e.g. Untermassfeld), and the Mid-

Pleistocene Homotherium latidens with new records recordedherein: Balve Cave in the Sauerland Karst, Zoolithen Cave in theFrankonian Karst, Ketsch Rhine River Terrace Upper Rhine Graben.The new Mid-Pleistocene H. latidens humeri are discussed alongwith the northern German Harz foreland Sch€oningen lake siteincomplete felid humerus, which is revised to a smaller-sized lionor female due to absence of the supracondylar ridge. The Germansaber-tooth catH. latidens records seem to reflect “dispersal events”within interglacials, several times in the Cromerian Complex MISduring the MIS 19 Interglacial (Mosbach, West Runton), the MIS 15and 11 as shown for Czech cave (Sluj IV) records, and in the MIS 9eHolsteinian Interglacial, (Sch€oningen). Probably, saber-tooth catsappeared again in the early Late Saalien two warm periods, theWackenian and Lietian Interstadials (MIS 7a, c), but they reachednot into the final Saalian (MIS 6) nor survived into the Late Pleis-tocene in Europe. No saber-tooth cat is proven from an archaeo-logical “killing” context. All known saber-tooth cat remains arefound in hyena den cave/open air sites or in carnivore conflictcontexts, including the Sch€oningen Lake site. Bite marks on a bovidtibia and lion humerus from the Sch€oningen site can be demon-strated to have the same hyena breaking scissor tooth impactmarkson their distal joints. Hyenas produced similar “diagonal cuts” atany guilt between joints for dismembering large body pieces (e.g.legs) to move those from the site to avoid further carnivore or evenhuman conflicts. Those bone damages were left most probably atSch€oningen and other Mid-Pleistocene sites by the early brownhyena Parahyaena brunnea mosbachensis (or spotted hyena Crocutacrocuta praespelaea/ultima), which is indirectly proven by its bitemarks, and coprolites, and possibly directly by a scapula.

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