Panthera Pardus ., 2003).Ouraim,However, Is Var

ICBS PROCEEDINGS and Merkenstein-cave(441m)nearBaden,Lower Austria the Wildkirchli-cave (1,420m)attheSäntis,Switzerland in Alpine bearcaves.However, tworecently revisedsites, INTRODUCTION Liguirië (Italië). Het luipaard bereikte slechtsincidenteelhet Alpengebied metuitzonderingvandezuidelijke Franse Alpen en lochgrot inZwitserland. Toch isdedistributieinhetOostelijk Alpengebied beperkt, vooral bijgrotere hoogtes. Pleistoceen bereikte hetluipaard ookhet Alpengebied. Dehoogstgelegenvondsten komen vandeDrachen Hundsheim (Oostenrijk), gevolgd doorresten uitdeRepolustgrot inStyria(Oostenrijk). Gedurende hetLaat- het Oostelijk Alpengebied enZwitserland. Hetoudstebewijsmateriaal komt uithetMidden-Pleistoceen van Dit artikel presenteert eenherzieningvandegevonden luipaardresten inde Alpen meteen specialefocus op Samenvatting exception ofthesouthernFrench Alps andLiguria, Italy. Alpine area islimited, especially athigherelevations. The leopard reached the Alpine area only occasionally with highest elevated findscomefrom theDrachenloch, Switzerland. Nonetheless, itsdistributionintotheEastern from Repolust-cave, Styria, Austria. DuringtheUpperPleistoceneleopard alsoreached thehigh Alps. The Switzerland. The oldestevidence comesfrom theMiddle PleistoceneofHundsheim, Austria, followed by remains The paper presents arevision ofleopard remains inthe Alps withspecialfocus ontheEastern Alpine area and Abstract OGY, GEOZENTRUM UZAII, ALTHANSTRASSE 14, A-1090 VIENNA, AUSTRIA, [email protected] GERNOT RABEDER OGY, GEOZENTRUM UZAII, ALTHANSTRASSE 14, A-1090 VIENNA, AUSTRIA, [email protected] MARTINA PACHER DURING THE UPPERPLEISTOCENE OF HUNTER THE ALPINEAREA RARE THE LEOPARD ( [ navicular bonewas found. Two remainsare currentlyhoused fragment mentioned byFreudenberg (1914)northefigured na confirmssixteen specimens.Neitherthethird metacarpal terial fromHundsheimattheNatural HistoryMuseum Vien 1914, Fig.XVIII:1r, Thenius, 1953). The revision ofthema ( lanx subsequentlyidentifiedas first evidenceofthecheetah listed andfiguredelevenremains amongthemadistalpha tioned inFreudenberg (1909).Lateron,Freudenberg (1914) come fromtheMiddlePleistoceneofHundsheim,firstmen REVISION OFSITES Switzerland. on arevisionofmaterialfromtheEastern Alpine areaand the Alps andtocompilemetricaldata.Specialfocusisgiven to summarizepossibleoccurrencesofPleistoceneleopardsin Diedrich, 2013;Hemmer sion ofPleistoceneleopards(eg.Hemmer&Schütt,1970; berg 1914)andlateronincludedinthetaxonomicdiscus been describedas from theMiddlePleistoceneofHundsheim,whichhavefirst published sofar. the opportunitytocompileandaudit Alpine leopardevidence identified skeletalelementsinthelattercave. Thisgivesus Pleistocene withtwenty-twointheformerrespectivelysix reveal morenumerousremainsoftherarehunter 42 Acinonyx intermedius Remains oftheleopard( The oldestremainsofleopardsintheEastern Alpine area The oldestknownremainsintheEastern Alpine areacome ]

Felis pardus UNIVERSITYOF VIENNA, INSTITUTEOFPALAEONTOLOGY, PALAEOBIOLOGY-VERTEBRATEPALAEONTOL UNIVERSITYOF VIENNA, INSTITUTEOFPALAEONTOLOGY, PALAEOBIOLOGY-VERTEBRATEPALAEONTOL n.sp.)atHundsheim(Freudenberg et al Panthera pardus ., 2003).Ouraim,however, is var. tulliana foss.(Freuden ) arerarefinds PANTHERA PARDUS ------assigned totheleopard (Fladerer, 1997b:). A patellafrom the phalanx (LMJ58278), originallyclassifiedaslynx, are er-cave, onecervicalvertebra (LMJ22648)andonemiddle finds, whoseidentificationisnot alwaysclear. AtHolzing ties totheleopardinsize(table8). was assignedtothelynx( confirmed. Anadditional proximalphalanx(LMJ55949) Mottl (1953)andoneproximalphalanx(LMJ57436)are one thirdmetacarpal(LMJ58280)listedasmetatarsalin only byasmallstream.Onepelvic-fragment(LMJ57435), mains fromBadl-caveoppositeofRepolust-cave,separated teeth (table4). B-14.0mm), dp4(L-14.1,B-5.0mm)andgermsofdeciduous mandible possessesthedc(L-13.6,B-9.5mm),dp3(L-10.8, juvenile (LMJ58978)isattributedto ever, onemt3(LMJ 57433)andamandiblefragmentof not befoundatthemuseum“Joanneum”inGraz.How and phalanges, soil arementioned(Mottl,1951:65-66,Fig.23). The canine a pelvicfragment(LMJ54612)fromtheauburnphosphate langes, onecanine,fourthmetacarpal(LMJ54613)and ditional faunalelements(Pacher, 2014). Two proximalpha towards theUpperPleistocenebasedonstratigraphyandad are mostlikelyfromthetransitionofMiddlePleistocene tions from440ma.s.l.(abovesealevel)to650m Pleistocene. All mentionedcavesiteslieatmoderateeleva older remains,aswellseveralfindsattributedtotheUpper Vienna (table2,table4-8,plate1). at thecollection Weinfurter atthe InstituteofPalaeontology Several cavesinthemountainarea nearGrazyieldsingle Probably ofsimilarchronologicalagearetheleopardre The leopardremainsfromRepolust-cavenearGraz,Styria The mountainareanearGrazyieldedchronologically

one withinventorynumberLMJ57434could Lynx lynx ), THE ) butshowscloseaffini Panthera pardus

. The ------CRANIUM

JUNI 2016 ICBS PROCEEDINGS AUTHORS PACHER MARTINA RABEDER GERNOT JUNI 2016

- , and the CRANIUM These finds were spelaea. Panthera spelaea var. . Felis pardus Felis pardus In the Austrian High Alps, the leopard is only listed at Austrian High In the Alpine area, the leopard is known from In the Southern The upper canine from Drachenloch is figured in Diedrich The upper canine from Drachenloch is figured Bächler (1936; 1940) ascribed the leopard remains from At Ranggiloch, the fragment of a phalanx is assigned Ochsenhalt-cave with a left distal fragment of an upper P4 The element was not available for 2007:72, Fig. 5). (Geyer, this study. (2007) mentioned a lower Toškan Divje babe I, Slovenia. m1 (length of paraconid = 9.8mm), a vertebra, a carpal bone, three medial phalanges and two distal phalanges from Panthera pardus table 19). New elements are labeled “WKV” for finds stud table 19). New elements are labeled “WKV” from Drachenloch are mentioned remains Two Vienna. ied in CXIII:1-2). and figured in Bächler (1940: 213 and plate The maxilla The canine is also labeled with “Drachenloch”. labeled The second metacarpal fragment was not available. comes also from the Drachenloch-cave and “AB 4-6 P.E.” was found within bear metapodials in the museum. The preserved Wildkirchli finds. (2013: fig. 6-5) among the (Diedrich total height of the incisors, 33mm and 36mm for 2013: fig. 6-3, 6-4), argues 2013: fig. 6-10) heavily abraded proximal radius (Diedrich shows affinities to a small-sized bear. to Wildkirchli the first evidence of the Pleistocene leopard in Switzerland. Nonetheless, the highest elevated occurrence lies at the Drachenloch (2,475m a.s.l.) followed by the Ranggiloch (1,845m a.s.l.) and the Chilchiloch (1,810m a.s.l.). In the Alpine area the highest elevated evidence comes Eastern Apart from from the Ochsenhalthöhle (1,660m a.s.l.), Styria. (1,477m a.s.l), remains of the leopard are scarce Wildkirchli Alpine sites and sometimes not well confirmed. in At 1964). to the leopard because of its dimensions (Koby, Chilchi-cave, the fragment of an upper P4 was as clearly identified as a right lower p3 from Schnurenloch (Koby, 1964:156, Fig. 31). CXI). Evidence of an upper M1 is based on Schmid (1940, CXI). Evidence of an upper M1 is based on - - et - - (table Panthera pardus is also mentioned at the Panthera pardus In course of this revision, twenty-one leopard remains are The most numerous remains at the high Alps are reported The most numerous remains at the high Undoubted leopard remains come from Merkenstein-cave, ., 2014) db = 34.7mm, btr = 24.4mm). already been figured by Bächler (1936: Fig.32, 1940: plate confirmed at Wildkirchli-cave (table 3); some of them have confirmed at large mammal remains were also examined. large Drachenloch, Wildkirchli and Wildenmannlisloch, financed Wildenmannlisloch, and Wildkirchli Drachenloch, by the Naturmuseum St. Gallen, Switzerland, the interesting from the Wildkirchli-cave, Switzerland. In course of a revi Wildkirchli-cave, from the Alpine sites sion of the cave bear remains from three Swiss exceed the values from the leopard of Cerè cave (Ghezzo al other Pleistocene elements at the site. Its dimensions (distal breadth (db) = 40.4mm, breadth of trochlea (btr) = 29.0mm) to the mentioned element of lynx (Wettstein & Mühlhofer, & Mühlhofer, to the mentioned element of lynx (Wettstein the from the preservation is different 1938). Surprisingly, 6), an upper incisor, and a right first tarsal bone. One distal 6), an upper incisor, humerus fragment from a leopard probably corresponds A recent revision yielded an additional pelvic-fragment, A for whose dimensions clearly argue of the p4 are on the lower range of values and are even small when compared to modern leopards (Nagel, 1997). stein & Mühlhofer, 1938:537, fig.5; Nagel, 1997). Both stein & Mühlhofer, The dimensions remains were not available for this study. its entrance at 441m a.s.l.. One lower p4 and a proximal phalanx have already been classified as leopard (Wett Lower Austria. This cave lies again at lower elevation with Austria. Lower 58909) labeled as male lynx is assigned to the leopard due to 58909) labeled as male lynx is assigned to the its size (table 8). not available at the collection of the museum “Joanneum” in not available at the collection of the museum (LMJ Graz. In addition, a middle phalanx from Spalten-cave phalanx (Fladerer & Frank, 1997) and listed for the Große phalanx (Fladerer & Frank, 1997) and listed The remains were 1997a). Peggauerwandhöhle (Fladerer, (Fladerer, 1997c). (Fladerer, based on a small-sized proximal Kugelstein-Tropfsteinhöhle, (Fladerer, 1997d:). The same holds true for a distal meta The same holds true for a 1997d:). (Fladerer, podial fragment (LMJ 58281) from the Fünffenstergrotte breadth of 18.7mm, a greatest length of 31.4mm and a depth breadth of 18.7mm, a greatest length of 31.4mm of leopards of 8.8mm the specimen falls within the range Luegloch (LMJ 58533) was referred to as lynx but the leop Luegloch (LMJ 58533) was referred to as lynx a greatest With ard is more likely because of its dimensions. Figure 1: Cave sites with Panthera pardus remains from the Eastern Alpine area and Switzerland (Abbreviations as in Table 1) Table as in Alpine area and Switzerland (Abbreviations remains from the Eastern pardus Cave sites with Panthera Figure 1: 1) Tabel en ZwitserlandAlpengebied (Afkortingen als in in het oostelijk pardus overblijfselen Grotten met Panthera Figuur 1:

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------Geologisch Rivista Mitteilungen , 342-348 Prähistorische CRANIUM L’Anthropologie 108 L’Anthropologie Quaternary Science , 375-391. , 320-325. 193-199. und pleistozäne Faunen

, 312-316. Buchdruckerei Tschudy & Co. Tschudy Buchdruckerei Pliozäne und pleistozäne Faunen Österreichs. und pleistozäne Faunen Österreichs.

, 311-312. , 308-310. und pleistozäne Faunen Österreichs. Mit und pleistozäne Faunen Österreichs.

Nouvelles Archives du Muséum de Histoire du Muséum de Histoire Archives Nouvelles Das Wildkirchli. in der Schweiz Das alpine Paläolithikum , 61-102. Pliozäne , 261-307. und pleistozäne Faunen Österreichs. Mitteillungen und pleistozäne Faunen Österreichs. Mitteillungen und pleistozäne Faunen Österreichs. , 197-222. Pliozäne

Felidae. , 167-193. Jahrbuch der kaiserlich-königlichen geologischen Reichs Mémoires du Muséum National d’Histoire naturelle n.s. sér. sér. n.s. naturelle du Muséum National d’Histoire Mémoires , 44-377. Pliozäne Pliozäne . Verlag Birkhäuser, Basel. Birkhäuser, Verlag . Fladerer, F. (1997a) Große Peggauerwandhöhle. in: Döppes, D., F. Fladerer, (1997b) Holzingerhöhle. in: Döppes, D., G. Rabeder F. Fladerer, in: Döppes, D., G. Rabeder (1997c) Fünffenstergrotte. F. Fladerer, (1997d) Luegloch bei Köflach. in: Döppes, D., G. F. Fladerer, am Kugelstein. in: Tropfsteinhöhle C. Frank (1997) F., Fladerer, (1909) Die Fauna von Hundsheim in Niederö W. Freudenberg, (1914) Die Säugetiere des älteren Quartärs W. Freudenberg, (2012) Der nächtliche Jäger als A., B. Horejs, B. Nessel Galik, Bächler, E. (1936) Bächler, E. (1940) Bächler, Pléistocène supérieur de la grotte Ballesio, R. (1980) Le gisement felids Pleistocene large The Middle R. Sardella (2012) Bona, F., Carnivores quaternaires du sud-est de la (1971) M.F. Bonifay, Pleistocene leopards across Europe – Diedrich, C.G. (2013) Late A. (2004) Étude taphonomique, paléoécologique et Echassoux, (Panthera pardus L., K.H. (2000) Ein Leoparden-Fund Fischer, , 221-227. G. Rabeder (Eds.) der Öster Mitteillungen der Kommission für Quartärforschung 10 der Wissenschaften Akademie reichischen (Eds.) Akade der Kommission für Quartärforschung der Österreichischen 10 mie der Wissenschaften (Eds.) Akade der Kommission für Quartärforschung der Österreichischen 10 mie der Wissenschaften Rabeder (Eds.) teillungen der Kommission für Quartärforschung der Österreichi 10 Akademie der Wissenschaften schen Döppes, D., G. Rabeder (Eds.) Mitteillungen der Kommission für Quartärforschung Österreichs. 10 Akademie der Wissenschaften der Österreichischen sterreich. anstalt LVIII(2) von Mitteleuropa, mit besonderer Berücksichtigung der Fauna in Niederösterreich nebst von Hundsheim und Deutsch-Altenburg Bemerkungen über verwandte Formen anderer Fundorte. 12(4-5) Abhandlungen N.F. Paläontologische Beute. Studien zur prähistorischen Leopardenjagd. Zeitschrift 87(2) St. Gallen. Die ältesten und Wildenmannisloch. Drachenloch im Wildkirchli, des Schweizer Altsteinzeit Niederlassungen aus der menschlichen landes France: Les Carnivores (Mammalia, de Jaurens a Nespouls, Corrèze, Carnivora) II. 18 de Lyon naturelle N-E Italy). di Soave (Verone, (Mammalia) from Brecce 118, e Stratigrafia Italiana di Paleontologia France. C, XXI population, highest elevated records northernmost European German complete skeleton in the Bosnia-Herzegowina Alps, in the Swiss Age cave art. Dinarids and comparison to Ice Reviews 76 de la second archéozoologique des faunes de grands mammifères (Roque Vallonnet moitié du Pléistocène inférieur de la grotte du Alpes-Maritimes, brune-Cap-Martin, France). 11-53. von Berlin und 1758) aus dem jungpleistozänen Rixdorfer-Horizont der Leoparden im Pleistozän Europas. Verbreitung die Reihe des Museum für Naturkunde in Berlin Geowissenschaftliche 3 REFERENCES ------Tri -

(Bona & Sar in Spain. The in Spain. ., 2012; Spassov Serbaro di Romag

., 2013) in addition to ., 2013) in addition to et al remains hamper a et al ., 2014) and Panthera fossilis Panthera pardus et al Panthera pardus

We would like to thank Ursula Göhlich and Alexander would like to thank Ursula Göhlich and We The leopard was a scarce element in the Eastern Alpine The leopard was a scarce element in the Eastern Based on literature, evidence stays scarce in the Italian in the stays scarce evidence on literature, Based 11: metatarsal 5, plantar view (NHMW 1909-II-174+175) plantar view metatarsal 5, 11: view (NHMW 1909-II-174); 8: proximal phalanx (NHMW 1909-II-171); 9a: metatarsal 4, plantar view (NHMW 1909-II-174); 9b: metatarsal 9b: (NHMW 1909-II-174); plantar view metatarsal 4, 9a: phalanx (NHMW 1909-II-171); proximal 8: (NHMW 1909-II-174); view (NHMW 1909-II-170); medial view tarsal 3, 10b: (NHMW 1909-II-170); view proximal tarsal 3, 10a: (NHMW 1909-II-174); view proximal 4, 1909-II-169); 4b: scaphoid, distal view (NHMW 1909-II-169); 5a: talus, dorsal view (NHMW 1909-II-176); 5b: talus, plantar view (NHMW plantar view talus, 5b: dorsal (NHMW 1909-II-176); view talus, 5a: (NHMW 1909-II-169); distal view scaphoid, 4b: 1909-II-169); proximal metatarsal 3, 7b: metatarsal (NHMW 1909-II-174); 3 plantar view 7a: dorsal (NHMW 1909-II-176); view calcaneus, 6: 1909-II-176); 1a: metacarpale 4, palmar view (IPUW, coll. Weinfurter); 1b: metacarpale 4, proximal view (IPUW, coll. Weinfurter); 2: middle phalanx (IPUW, coll. coll. middle phalanx (IPUW, 2: Weinfurter); coll. (IPUW, view proximal metacarpale 4, 1b: Weinfurter); coll. (IPUW, palmar view metacarpale 4, 1a: (NHMW view proximal scaphoid, 4a: (NHMW 1909-II-167); lingual view P3sup, 3b: (NHMW 1909-II-167); buccal view P3sup, 3a: Weinfurter); Museum of Vienna). All photos lifesize pictured All photos lifesize Vienna). Museum of afgebeeld levensgroot Alle foto’s pardus uit Hundsheim. Panthera Gesecteerde elementen van Plaat 1: Plate 1: Selected elements of Panthera pardus from Hundsheim (IPUW Institute of Palaeontology, University of Vienna, NHMW Natural History NHMW Natural Vienna, University of pardus from Hundsheim (IPUW Institute of Palaeontology, Selected elements of Panthera Plate 1: Gallen, Switzerland and the project „Lunzer Bärenhöhlen“, Kultur Niederösterreich. Vienna and the Wasserkluster Lunz kindly supported our Wasserkluster and the Vienna The study is financed by the Naturmuseum St. work as well. anneum Graz for their friendly help and providing the leop Paleontology, of Institute The sites. various from material ard Bibl, Natural History Museum in Vienna, Toni Bürgin, Bürgin, Toni Vienna, Bibl, Natural History Museum in Naturmuseum St. Gallen, Martin Gross, Landesmuseum Jo ACKNOWLEDGMENTS & Raychev, 1997). & Raychev, on unclear dated finds only (eg. Galik Pleistocene refuge area for discussed Holocene survival of leopards in Europe is based its occurrence during MIS 3 (Ghezzo & Rook, 2015), its occurrence during MIS 3 (Ghezzo & Rook, an Upper while Sauqué & Cuenca-Bescós (2013) suggest BP (Rabeder & Symeonidis, 1995) and a sample from Equi, BP and hence confirms Alps dated at 33,000+/-260 BP Apuan out Europe. A direct date on leopard bones from Vraona gave Vraona direct date on leopard bones from A out Europe. and 24,490+/-1350 a conventional age of 17,805+/-1000 BP radiocarbon dates on through detailed time resolution of the leopard distribution the Alpine area and more northern regions only occasion the the nearly complete absence of direct Unfortunately, ally. & Rook, 2015), it seems that the Pleistocene leopard favored & Rook, 2015), it seems that the Pleistocene and migrated into moderate climatic conditions through time France (Ballesio, 1980), sites in Spain (see Sauqué & France (Ballesio, 1980), sites in Spain (see Italy (Ghezzo Cuenca-Bescós, 2013) and especially at Equi, southern Europe, such as Vraona, Greece (Nagel, 1999), Greece (Nagel, Vraona, southern Europe, such as 1997), Jaurens, agalnata cave, Bulgaria (Spassov & Raychev, ter living conditions at the southern fringe of the Alps. Given the ter living conditions at the southern fringe of of remains in the number of sites with a considerable number ably reached higher elevated sites such as Wildkirchli and Wildkirchli ably reached higher elevated sites such as found bet whereas leopards Drachenloch only occasionally, area and Switzerland during the Upper Pleistocene. It prob area and Switzerland during the Upper Pleistocene. CONCLUSION sites (eg. Valensi & Pasthi, 2004; Bonifay, 1971). 2004; Bonifay, & Pasthi, Valensi sites (eg. starting from Early Pleistocene remains at Vallonnet (Echas Vallonnet remains at starting from Early Pleistocene Middle to Upper Pleistocene soux, 2004) towards numerous sakis & Palombo, 1979). Leopards get considerably more sakis & Palombo, 1979). Alps Alps and the southern French abundant in the Ligurian yielded two remains (Tagliacozzo yielded two remains (Tagliacozzo 1957) and Grotta di Sambughetto (Kot Zandobbio (Vialli, attributed to a lion-like cat, attributed to a lion-like di Fumane The Upper Pleistocene from Grotta della, 2012). nano (Kotsakis & Palombo, 1979). The proposed evidence at The proposed 1979). nano (Kotsakis & Palombo, have recently been and Soave-Sentiere Tenda Soave-Monte research leopards are confirmed at the Middle Pleistocene at the Middle Pleistocene are confirmed research leopards from Cerè cave (Ghezzo listed in Diedrich (2013) and five in Sauqué & Cuenca-Bes (2013) and five listed in Diedrich compilations and own Following the previous cós (2013). limited number of sites in this area with one site mentioned with one site mentioned of sites in this area limited number Three localities are (1940) and Fischer (2000). by Schmid Alps and is mainly restricted to sites at the southern Alpine Alpine the southern to sites at restricted is mainly Alps and a European finds show compilations of fringe. Previous

] 44 [ ICBS PROCEEDINGS ihre eiszeitlichenBewohner(miteinemBeitragvon V. Maurin). logica Romana18 (L.) delPleistocenemediosuperioredeMonteSacro(Roma). Bernensia III D., W. Flükiger, A. Andrist (Eds.) Panthera). masse ältest-undaltpleistozänereuropäischerPantherkatzen(genus 229 ars. zur Kenntnisder Variabilität und Verbreitungsgeschichte desJagu Mosbach-Sanden (Wiesbaden, Hessen,Deutschland).EinBeitrag gambaszoegensis (Kretzoi1938)ausdenfrühmittelpleistozänen Northeastern Italy). lerian Canidae,FelidaeandMustelidaeoftheCerèCave(Verona, 131-151. morphology, andpaleoecology. (Felidae, Mammalia)recordfromEqui(Massa,Italy):taphonomy, projekt. [ Tabel 1: Lijstvan grottenmetPanthera pardusresteninhetalpiene gebied Table 1: Listofcavesiteswith Panthera pardus remainsfromthe Alpine area 46 Mottl, M.(1951)DieRepolusthöhlebeiPeggau(Steiermark)und Kotsakis, T., M.R.Palombo(1979)UncraniodiPantherapardus Koby, F. E.(1964)Die Tierreste derdreiBärenhöhlen.in: Andrist, Hemmer, H.,G.Schütt(1970)Körpergrösse undExtremitäten Hemmer, H.,R.D.Kahlke, T. Keller(2003)Pantheraonca Ghezzo, E.,D.F. Berté,B.Sala(2014) The revaluationofGa Ghezzo, E.,L.Rook(2015) The remarkable Geyer, E.(2007)10JahreForschungenim Weißenbach-Höhlen ] , 31-60.

Sa Fu GPW DL DB Ch Ce Ba Abbrev. Zd Wk TH Sp Ro Sn Rep Ra OH Mst Lu Ff HH Hz Neues JahrbuchfürGeologieundPaläontologie Abhandlungen Die Höhle59(1-4) Mainzer naturwissenschaftliches Archiv 9 , 149-160. , 137-155. Zandobbio Wildkirchli Tropfsteinhöhle Spaltenhöhle Serbaro diRomagnano Schnurenloch Repolusthöhle Ranggiloch Ochsenhalthöhle Merkenstein Luegloch Fünffenstergrotte Hundsheim Holzingerhöhle Grotta diSambughetto Grotta diFumane Gr. Peggauerwandhöhle Drachenloch Divje babe Chilchihöhle Grotte delCerè Große Badlhöhle Site Quaternary International339-340 , 69-73. Quaternary ScienceReviews110 Das SimmentalzurSteinzeit. Acta Panthera pardus , 132-146. 750m 495m A.s.l 370m 1,477m 482m ~490m 1,230m 525m 1,845m 1,660m 441m 550m 441m 270m 650m 350m 510m 2,445m 450m 1,810m , 76-89. Geo -

, - - - -

MP MP/UP Chronology MP MP UP UP UP MP UP MP/UP UP UP UP UP UP MP UP UP UP UP UP UP UP Zeitschrift fürSäugetierkunde15 Gebiss pleistozänerundrezenterLeopardenandererFeliden. the UpperPleistocene. last EuropeanRefugiumofPantherapardusLinnaeus1758during Helléniques 36 „Vraona“ auf Attica inGriechenland. eonidis, N.,G.Rabeder(Eds.) der Höhlensedimentevon Vraona (Attika,Griechenland).in:Sym Untersuchungen. (Steiermark, Österreich)–ErsteErgebnisse derpaläontologischen Geologie undPaläontologie,Monatshefte3 leopard fromthePleistoceneof Vraona/ Greece. 224. Mitteilungen desNiederösterreichischen Landesmuseum aus derHöhlevonMerkenstein/Niederösterreich. Technik amLandesmuseumJoanneum11 Steiermark. Murban (Eds)EiszeitforschungendesJoanneumsinHöhlender Archaeologica8, Austriaca Schmid, E.(1940) Variationsstatistische Untersuchungenam Sauqué, V., G.Cuenca-Bescós(2013) The IberianPeninsula,the Rabeder, G.,N.Symeonidis(1995)StratigraphieundChronologie Pacher, M.(2014)Dieeiszeitliche Tierwelt derRepolusthöhle Nagel, D.(1999)Pantherapardusvraonensisn.ssp.,anew Nagel, D.(1997)Pantherapardusundspelea(Felidae) Mottl, M.(1953)DieErforschungderHöhlen.in:M.,K. Mitteilungen desMuseumsfürBergbau, Geologieund , 107-109. Schild vonSteier26,2013/2014 5 3+1? NISP 1 22 1? 1? 1 5+1? 1 1 6 1? 1? 17 2? 2 2? 3 8 1 Quaternaire 24(1) 1-78. Das JungpleistozäninderHöhlevon Ghezzo etal.,2014 Mottl, 1953,owndata References Vialli, 1957 Schmid, 1940,owndata Fladerer &Frank,1997 own data Kotsakis &Palombo,1979 Koby, 1964 Mottl, 1951,owndata Koby, 1964 Geyer, 2007 Nagel, 1997,owndata Fladerer, 1997d Fladerer, 1997c own data Fladerer, 1997b Kotsakis &Palombo,1979 Tagliacozzo etal.,2013 Fladerer, 1997a Bächler, 1940,owndata Toškan, 2007 Koby, 1964 , 1-179. Annales GéologiquesdePays , 14-58. , 35-48. , 129-150. Neues Jahrbuchfür , 238-254. Wissenschaftliche

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, 514-558. Divje babe Divje babe Atti della Società CRANIUM , 256-272. Archiv für Naturgeschichte N.F. 7(4) N.F. Naturgeschichte für Archiv table XVIII:3h-i table XVIII:3h-i table XVIII:3h-i table XVIII:3f table XVIII:3e Freudenberg (1914) Freudenberg table XVIII:2c-d table XVIII:3b table XVIII:3a table XIX:F , 51-79. Valensi, P., E. Psathi (2004) Faunal Exploitation during the E. Psathi (2004) Faunal Exploitation P., Valensi, vertebrati della breccia ossifera dell’inter (1957) I V. Vialli, (1938). Die Fauna der Höhle von Mühlhofer F. O.V., Wettstein, Toškan, B. (2007) Remains of Large Mammals from Divje babe from Divje Mammals of Large (2007) Remains B. Toškan, I Upper Pleistocene Palaeolithic site in Slovenia. Part I: Geology in Slovenia. Part I: Geology Palaeolithic site I Upper Pleistocene Sloveniae Archaeologici Institutti Opera and Palaeontology. 221-278. Italy France and North-western in South-eastern Middle Palaeolithic 14 Journal of Osteoarchaeology International (Bergamo). glaciale Riss-Würm di Zandobbio e del Museo Civico di Storia naturale in Italiana di Scienze naturali Milano XCVI Merkenstein in N.Ö. I: stratigraphy, taxonomy and biometry. in: Turk, I. (Ed.) I. Turk, in: and biometry. taxonomy I: stratigraphy, - - - - as Machairodus as medial phalanx rearticulated articulated articulated articulated Comment Zooar 1909-II-170 1909-II-171 1909-II-172-1 1909-II-168 Weinfurter IPUW-coll. Weinfurter IPUW-coll. 1909-II-174+175 1909-II-174 1909-II-169 1909-II-169 1909-II-176 1909-II-176 1909-II-176 1909-II-170 Inventory number 1909-II-167 1909-II-174 1909-II-174 1909-II-174 , 71-96. Neues Jahrbuch für Geologie und Paläon Neues Jahrbuch für Geologie , 225-238. Springer Science & Business Media, Dordrecht, & Business Media, Springer Science . Middle phalanx Mc4 sin Caudal vertebra Proximal phalanx Proximal phalanx Tarsal III sin Tarsal III dex Tarsal Calcaneus sin Naviculare sin Astragalus dex Astragalus sin Metapodial distal Scapholunatum dex Mt4 sin Mt5 dex Mt2 dex proximal Mt3 dex Element P3 sup Spassov, N., D. Raychev (1997) Late Wurm Panthera pardus Wurm N., D. Raychev (1997) Late Spassov, Thenius, E. (1953) Gepardreste aus dem Altquartär von Hunds Gepardreste aus dem Thenius, E. (1953) Tagliacozzo, A., M. Ronadini, I. Fiore, M. Gala, M. Peresani M. Gala, M. I. Fiore, A., M. Ronadini, Tagliacozzo, Tabel 2: Gereviseerde lijst van luipaardvondsten uit Hundsheim lijst van Gereviseerde 2: Tabel Table 2: Revised list of leopard remains from Hundsheim Revised 2: Table Historia naturalis bulgarica 7 Historia naturalis bulgarica remains from Bulgaria: the European leopards and the question of remains from Bulgaria: the until the Holocene on the Balkans. the probable species survival tologie Monatshefte 5 heim in Niederösterreich. heim in Niederösterreich. anthropologie 129-150. chaeology and Modern Human Origins: Human Hunting Behavior Human Hunting Behavior Modern Human Origins: chaeology and and Paleo Paleobiology Pleistocene, Vertebrate during the Later (2013) Animal Exploitation Strategies during the Uluzzian at Grotta the Uluzzian during Strategies Exploitation Animal (2013) J.D. Speth (Eds.) Italy). in: Clark, J.L., di Fumane (Verona,

] 46 [ ICBS PROCEEDINGS [ Tabel 3: Gereviseerde lijstvan Panthera pardusvondstenuitDrachenloch (DL)en (Wk) Wildkirchli Table 3: Revised listofPanthera pardusfindsfromDrachenloch (DL)andWildkirchli (Wk) 48 ]

Wk-18 Wk-17 Wk-16 Wk-15 Wk-14 Wk-13 Wk-12 Wk-11 Wk-10 Wk-9 Wk-8 Wk-7 Wk-6 Wk-5 Wk-4 Wk-3 Wk-2 Wk-1 DL DL DL Material Wk-21 Wk-20 Wk-19 --- WKV-37 WKV-341 WKV-70 WKV-59 I 2,3m I 2,30m K 1m K 3m AIII 1,80m A4 1,3m K 1,5m K 3m K 2,60m I 2,10m --- K 2,9m WH AI 2,3m WH m2401,6m W WH m2332,7m K 2,9m AB4-6 P.E HIII 5,5Sch.41m?(80cm)4.5.20 H3 P512/IX/19214S.. Labeled calcaneus calcaneus astragalus mc5 mc4 ulna thor.-vertebra P4 sup C sup C sup C sup maxilla+p3+p4 mc2 maxilla+P4 Csup Element M1 sup ph2 ph2 ph1 postIV P3 sup ph3 ph1 postIV ph1antIV mt5 sin dex dex dex dex dex sin dex dex sin dex sin sin dex Side ? dex dex Taf. CXI Taf. CXI Taf. CXI Taf. CXI Taf. CXI Taf. CXI Taf. CXI Taf. CXI Taf. CXIII Taf. CXIII Bächler (1940) Schmid (1940) Taf. CXI CRANIUM

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- After Ghez zo et al., 2014: 84 dd 13.3 13.4 14.7 15.0 CRANIUM 13.0 dcb 12.6 13.2 14.4 15.0 14.6 12.8 13.9 14.9 13.7 db 14.5 13.3 14.5 15.2 16.3 16.2 16.1 14.8 14.8 corroded table 19 + own Schmid, 1940: + own Schmid, 1940: table 19 Schmid, 1940: table 19 Schmid, 1940: table 19 Schmid, 1940: table 19 Schmid, 1940: table 19 1940: plate CXIII after Bächler, Nagel, 1997: 216, table 1 1964:155 Koby, germ Comment 9.5 9.9 SD 9.3 9.0 9.6 10.8 10.1 11.8 11.8 10.8 8.9 9.1 9.0 8.1 9.4 Bdi pd 16.6 17.6 17.2 17.9 18.5 23.0 21.0 19.0 -- 9,6 9,6 7.4 9.2 Bm 15.0 18.0 15.9 15.0 16.1 17.0 13.5 12.0 14.8 14.0 17.8 pb 14.8 8.7 7.5 13.5 13.6 -- Bme 8.8 ------4.4 -- 8.8 B 10.8 -- 10.4 10.7 94.4 84.8 96.0 99.0 96.6 62.6 66.9 75.0* 76.0 69.7 gl 70.5 19.4 19.0 18.0 18.7 25.9 25.8 7.8 25.0 15.6 L 13.8 -- -- 13.7 Mt2 Mt3 Mt3 Mt4 Mt5 Mt5 Mc2 Mc2 Mc4 Mc4 Mc4 Mc5 Elem Mc3 LMJ 57433 1909-II-174 1909-II-175 1909-II-174 1909-II-174 AB4-6 P.E. IPUW LMJ 54613 Inv. LMJ 58280 V.5108-1 P4 sup (Max.) p4 inf (?) p3 inf m1 inf P4 sup (Max.) P4 sup sin M1 sup C sup dex C sup sin P3 sup P3 sup (Max.) Element C sup dex C sup dex Rep Mst Sn Wk-21 DL Wk-5 Wk-1 Wk-1 Wk-2 HH Wk-3 Wk-4 Site DL Wk-13 HH HH HH Rep Wk-9 HH Rep Wk-8 HH DL Ba Ce Site Tabel 5: Maten van metapoden van Panthera pardus van de greviseerde sites (gl = grootste lengte, pb = proximale breedte, pd = proximale pd = proximale breedte, pb = proximale sites (gl = grootste lengte, de greviseerde pardus van Panthera metapoden van Maten van 5: Tabel *gereconstrueerd dd = distale diepte), de condyle, dcb = distale breedte van db = distale breedte, SD = kleinste diafysebreedte, diepte, Table 5: Measurements of metapodials from Panthera pardus from the revised sites (gl = greatest length, pb = proximal breadth, pd = breadth, pb = proximal sites (gl = greatest length, pardus from the revised Measurements of metapodials from Panthera 5: Table *reconstructed dd = distal depth), dcb = distal condyle breadth, db = distal breadth, SD = smallest diaphyseal breadth, depth, proximal Tabel 4: Maten van tanden van Panthera pardus van de gereviseerde sites (L= lengte; B = breedte; Bme = mesiale breedte; Bm = breedte over breedte over Bm = breedte; Bme = mesiale B = breedte; sites (L= lengte; de gereviseerde pardus van Panthera tanden van Maten van 4: Tabel Bdi = distale breedte) de tand; het midden van Table 4: Measurements of teeth from Panthera pardus from the revised sites (L = length, B = breadth, Bme = mesial breadth, Bm = breadth in Bm Bme = mesial breadth, B = breadth, sites (L = length, pardus from the revised Measurements of teeth from Panthera 4: Table Bdi = distal breadth) the middle of the tooth,

] 48 [ ICBS PROCEEDINGS breedte van articulatievlak) Tabel 7: elementenvan Matenvan Panthera overige pardusvan degereviseerde sites(gb=grootstebreedte, gl=grootstelengte, baf= surface) breadth ofarticular Table 7: MeasurementsofadditionalelementsfromPanthera pardusfromtherevised sites(gb=greatestbreadth, gl=greatestlength, baf= rand, SH=kleinstehoogtevan iliumshacht, SB=kleinstebreedtevan iliumshacht, SC=kleinsteomtrekvan iliumshacht) Tabel 6: Matenvan hetbekken van Panthera pardusvan degereviseerde sites(LA=lengtevan acetabulum, LAR=lengtevan acetabulumop rim, SH=smallestheightofiliumshaft, SB=smallestbreadthofiliumshaft, SC=smallestcircumference atiliumshaft) Table 6: MeasurementofpelvicbonesfromPanthera pardusfromtherevised sites(LA=lengthofacetabulum, LAR=lengthofacetabulumon [ diepte, SD= kleinstebreedtevan diafyse, db=distalebreedte) *aanvankelijk alslynx gedetermineerd Tabel 8: Matenvan falangen van Panthera pardusvan degereviseerde sites(gl =grootstelengte, pb=proximale breedte, pd=proximale depth, SD=smallestdiaphysealbreadth, identifiedaslynx db=distalbreadth)*originally Table 8: Measurements ofphalangesfromPanthera pardusfromrevised sites(gl=greatestlength, pb=proximal breadth, pd=proximal 50 Site Wk-20 Wk-19 DB DB DB Sp* Hz* HH Ce Wk-18 Wk-15 Wk-14 TH Rep Mst HH HH Ba* Ba Site ] Site Rep Mst Ba

HH HH HH HH HH HH Inv. 1909-II-170 1909-II-170 1909-II-176 1909-II-176 1909-II-176 1909-II-169 V.5078-50 WKV-70 LMJ 57434 -- 172-1 171 LMJ 55949 LMJ 57436 Inv. WKV-37 WKV-341 LMJ 58909 LMJ 58279 IPUW Inv. LMJ 54612 NHMW-GP LMJ 57435 Ph2 Ph2 Ph2 Ph2 Ph2 Ph2 Ph2 Ph2 Ph2 Ph1 post Ph1 post Ph1 ant Ph1 ant Ph1 post Ph1 Ph1 Ph1 Ph1 Ph1 Element Element tarsal 3 tarsal 3 Navicular Calcaneus Astragalus Scapholun. LA 26.0 32.3 29.0 gl 23.0 31.9 31.6 30.5 27.5 29.0 (20.8) 26.0 30.3 38.5 40.4 45.5 16.1 41.0 41.4 39.4 39.3 38.0 42.7 gb 16.0 13.6 29.9 33.5 31.6 27.3 LAR pb 11.1 13.5 10.7 (9.0) 12.1 11.4 15.3 16.7 16.3 10.1 15.0 13.4 14,5 13.9 16.3 31.4 41.5 gl 29.5 28.1 26.0 69.2 39.0 24.7 pd 17.0 9.6 9.5 12.7 11.5 13.2 12.3 12.5 SH 37.0 25.1 35.3 SD 7.1 7.2 5.5 5.4 8.4 7.3 9.5 9.8 10.3 8.3 10.0 10.0 10.7 12.0 baf 29.2 23.5 26.5 db 11.0 12.5 13.0 11.3 10.7 8.6 8.0 11.0 10.9 11.9 13.6 12.8 8.5 10.3 11.9 11.8 12.1 depth 14.4 13.9 10.0 33.0 23.3 16.4 SB 17.0 9.5 17.7 After own data own data Toškan, 2007:231,Fig.11-12 Toškan, 2007:231,Fig.11-12 Toškan, 2007:231,Fig.11-12 own data juvenile, owndata own data Ghezzo etal.,2014 own data own data own data Fladerer &Frank,1997 Mottl, 1951:66 Nagel, 1997:218,table2 own data own data own data own data After own data own data Freudenberg, 1914:191 own data own data own data SC 64.0 90.0 98.0 CRANIUM

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