ICBS PROCEEDINGS chronological positionbytheassumptionthatinterglacial „Drachenhöhle“ ofMixnitz.MariaMottl(1967)specifiedthe these cavebearslivedatthesametimeasof spelaeus deningeroides mountain area(Styria, Austria) wasdescribedas primeval bearremainsoftheRepolustcaveinGrazer literature byMariaMottl(1964). The relativelysmalland TYPE LOCALITY scrofa men vandeoverblijfselen vanstekelvarken ( vastgelegd.soort Overeenkomsten metdetypelocaliteitRepolustgrot kunnenworden gezieninhetvoorko fossiele faunauiteenkleinegrot inhetzuidenvan Wenen. Hiermee isdederde vindplaatsvandezemysterieuze De kleineenprimitieve grottenberensoort Samenvatting implies thesamegeologicalagefor bothfaunas. of porcupine ( enigmatic species. SimilaritiestothetypelocalityRepolustcave canbeseenintheappearance oftheremains of thefossil faunaremains locatedinasmallcave inthesouthof Vienna. This establishedthethird siteofthis The smallandprimitive cave bearspecies Abstract [email protected] MARTINA PACHER [email protected] CHRISTINE FRISCHAUF [email protected] GERNOT RABEDER DENINGEROIDES A NEWREFERENCEOF [ meters. jama arelocated attherelativelylowaltitudeof about 520 (Rabeder cave bearsisinterpretedasanadaption tothelifeinthisarea areunexplained. A reductionin sizeofhigh Alpine bach orHundsheimthesmalldimensions oftheRepolust- deningeroides both Urside-populationsareassociatedtothespecies taxonomical positionforbothfaunasisassumed. Therefore in evolutionarylevelanddimensionsomuch,thatthesame covered, whichresembledthematerialofRepolustcave merous remainsofasmallandprimevalcavebearwerere “Herkova jama“closetoRadljeobDrava(Slovenia).Nu Pohar THE SECONDSITE bears. cf. companying fauna(forexample correlated tothelateMiddlePleistocene,becauseofac thermore theyhypothesizedthatthefaunaofthiscaveis remains oftheRepolustcaveto and )indicatestheRiss/Würm-interglacial. character oftheaccompanyingfauna(especiallyporcupine 8 ] The taxonof In comparisontotheclassicalDeningeri-bears fromMos In theyears1977-1979and2001-2005(Pohar1981, Later revisors(Döppes&Rabeder, 1997)positionedthe

mosbachensis . diteengelijke Ookimpliceert geologischeleeftijdvoor beidefauna’s. et al. et al 2003)anexcavationtookplaceinthesmallcave . . 2008)buttheRepolust caveandtheHerkova deningeroides Hystrix vinogradoviHystrix ) andthelowevolutionarylevelofcave INSTITUTEOFPALAEONTOLOGY, UNIVERSITYOF VIENNA, ALTHANSTR. 14, 1090 VIENNA, AUSTRIA, INSTITUTEOFPALAEONTOLOGY, UNIVERSITYOF VIENNA, ALTHANSTR. 14, 1090 VIENNA, AUSTRIA, n.ssp. INSTITUTE OFPALAEONTOLOGY, UNIVERSITYOF VIENNA, ALTHANSTR. 14, 1090 VIENNA, AUSTRIA,

based ontheassumptionthat . Canis mosbachensis,Equus ), thelarge-sizedhorse was establishedinthe INLOWER AUSTRIA Ursus deningeroidesUrsus Ursus deningeroidesUrsus Ursus Hystrix vinogradoviHystrix Fur - Ursus - - - - Equus in themunicipality of Ternitz (districtNeunkirchen, Lower cave faunasincluding thefaunaofsmallFlatzer cave trian government istherevisionofseveralLower Austrian of Flatz). The objectofaproject fundedbytheLower Aus fossil faunaofthe“Flatzer Tropfsteinhöhle excavation butdetectedwithina revisionofthelongknown remains of SITE THE THIRD thetically theyareeligibleasanancestorofthese. and thereforenotpartofthe the faunas oftheRepolustcaveandHerkovajama. lower levelin Azé 1-3thanin thetypically lust andHerkovafauna. (ca. 4-5%)biggerthantheequivalentelementsofRepo and theHerkovajama(Frischauf of the 2011). Followingdifferences aroseafteradirectcomparison Azé 1-3mustbeolderthan190.000years(Barriquand corridor betweenthesite Azé 1-2and Azé 1-3, thefaunaof the Uranium-Thoriumdatingofsinterthatsealed the smalldimensionsandprimevalcharacter. Dueto bear-fauna of Azé 1-3as partly separated by sinter “curtains”. Argant (1991) described ing. The sites Azé 1-1to1-4arelocatedinacorridorand Mâcon (Burgundy, France)thebearremainsaredominat FRANCE) THE GROTTE OF AZÉ 1 (BURGUNDY, The detectionofthethirdsitewas asurprisebecausethe The cavebearremainsfrom Azé 1-3aretobeassigned The evolutionarylevelofthe4 The teethandmetapodialbonesof Azé 1-3areexplicitly In thefossilfaunasoffamousGrotted`Azé1near URSUS URSUS U. deningeri Mottl 1964wasdeterminedduringtherevision ), hetgrote paard Mottl1964isgevonden tijdensderevisie vande cf. Ursus deningeroides- mosbachensis Ursus deningeroides group. They arebiggerandmoreprimeval and Equus Ursus deningeroides faunas fromtheRepolustcave Sus scrofa. U. deningeroides cf. were not th et al premolarsisofamuch mosbachensis . 2013): Furthermore this

recovered atanew ˮ deningeroides

(Dripstone cave taxabuthypo because of en Sus et al - - - - - . - CRANIUM

JUNI 2016 ICBS PROCEEDINGS AUTHORS RABEDER GERNOT CHRISTINE FRISCHAUF PACHER MARTINA

- JUNI 2016 -

- - - (mor Ursus group . 2004. U. denin et al and group, because CRANIUM U. spelaeus Rabeder U. deningeroides U. deningeroides U. spelaeus- Ursus deningeri Ursus ingressus, U spealeus eremus Ursus ingressus, Ursus ingressus The premolars show only primeval from the sites Gamssulzen cave, Schwa of the Repolust and Herkova cave. The most The most of the Repolust and Herkova cave. ladinicus

The differences of the third lower incisors are The differences the ursid material of the Flatzer cave contains three the ursid material of the Flatzer cave contains whereas they are absent at the Ursus P4 inf. + P4 sup.: Peculiar is the relatively long but small shape of the two The third i2 inf. (Fl-N37) is bigger and has a pronounced The third i2 inf. (Fl-N37) is bigger and has Because the i2 inf. of i3 inf: Premolars The two mandibles where the bone structure between the Following elements are assigned to the taxa because of the to the taxa because of elements are assigned Following branches, 1 mandible 1 mandible with both 1 maxillare, of the Flatzer cave can be dated The fauna teeth are The auburn and black spotted mandibles and i2 inf: fragment an alveole for the p3 cannot be identified clearly. The p4 inf.-type B1 (with paraconid but morphotypes. without metaconid) appears only very seldom at the typically The same applies for morphotype “A” and “A/B” cave bears. on the upper premolars. p4 inf. But they might originate from one individual. because they are dominating geroides (Rabeder 1999: 78, tab. 50). (spelaeus), see mesoconid what is equal to morphotype “s” Fig. 2. and “s” morphotypes, benreith cave and Conturines cave are only i2 inf. from the morphology ensures that the two smaller Flatz belong to the “deningeri-form”. The lingual surface of typical cave bears similar prominent. is structured by vertically running enamel edges (mesial and distal edge) and furrows (sulcus mesialis and s. distalis) (morphotype C and D) whereas these features are low developed or missing on the teeth of The three smaller i3 inf. of the Flatzer and B). A photype and B) correspond with the level of the A cave (morphotype The bigger i3 inf. (inv.nr. Repolust and Herkova jama cave. Fl-10) complies with the typical it belongs to the C morphotype (see Fig. 3). canine and the p4 is present, have no alveole for another On the maxillary premolar so the dentition formula is 3113. dimension, morphology and the conservation status of the conservation status of morphology and the dimension, enamel: bones and the 5 canines, 4 premo fragment, 14 incisors, dex, 1 mandible bones, 1 metapodial bone, 9 lars, 14 molars, 4 autopodial phalanges from the The bones periods. back to two clearly different characteristic-yellow to auburn older period are coloured encrustations and are more heavy with brownish and black recognized in the heavier specific fossilized which can be suggest that they are younger. weight. Some of the remains to light brown and are without the au They are shaded gray furthermore they are clearly bigger and burn and black spots, most likely belong to teeth show a of consistently smaller dimensions and the for primeval character (s. tab. 1-4) that is also typical deningeroides to the typical cave bears can be detected obvious differences in the incisors. cusp, named “mesoconid” The mesial well preserved i2 inf. on two of them (Rabeder 1999) is only low or not developed The assigned morphotype is defined as (Fl-N32 and Fl-N34). “d” (deningeri) “d/s” (deningeri/spelaeus transition form) or MATERIAL - - - - possi , ˮ Spelaeus (2003) cfr. cfr. . were also found (Mottl, “Ursus Vulpes, Martes, Meles, Cervus, Vulpes, furthermore deer and bison. Hystrix Marmota Mottl, 1964 and also existed in the Flatzer fauna. The also existed in the Flatzer fauna. U. deningeri)

1997 Ursus deningeri Döppes & Rabeder (1997) 2003 Ursus deningeroides Pohar & al. Ursus deningeroides Ursus deningeroides 1964 Ursus spelaeus deningeroides n.ssp. Mottl (1964) It was a big surprise when we found a mandible of a unit 3: heavy and strong fossilized bones with gnawing unit 2: Late Pleistocene bones and teeth of the typical cave unit 1: Holocene remains of wild and domestic unit 1: Holocene remains of wild and domestic Although there has been no information about the find spot find the about information no been has there Although The first processing of the fauna was made by E. Thenius by E. The first processing of the fauna was made SYSTEMATIC DESCRIPTION SYSTEMATIC and black spots) confirmed all suspicions. of the other remains with the same characteristi cally conservation (relatively heavy fossilized with auburn comparison of the dimensions and morphology of the man dibles, teeth and metapodial bones as well as the comparison 1964, 1967). Instantly suspicion was raised that the small Ursus deningeroides traces. Thus an analogy can be seen in the fauna of the Repo traces. lust cave where remains of Neunkirchen. Furthermore the conspicuous gnawing traces, Thenius already noticed, could be identified as life which porcupine within the material of the municipal museum of bly related to marks: a primitive cave bear ( Capreolus, Bison, Bos Capreolus, bear and cave lion as well as chronological fauna levels by examining the conservation chronological fauna levels by examining the status of the teeth and bones: of the fossils, Thenius was able to separate three different Thenius was able to separate three different of the fossils, Neunkirchen. at the Lower Austrian State Museum of St. Pölten. New at the Lower the material of insights could be gained by later studies on (1949) and was based on the material that has been stored (1949) and was based on the material that has and the municipal museum of Neunkirchen. Austria, see Fig.1). The fossils of this cave are stored in two Austria, see Fig.1). Austrian State Museum of St. Pölten collection: the Lower Bergfreunde“). Er is geen informatie over de volgorde van de lagen. de Er is geen informatie de volgorde van over Bergfreunde“). Geologie: Steinalm kalksteen (Midden-Trias) Geologie: lokale groep mensen („Die 1904 tot 1906 door een Opgravingen: grottenregistratienr. 1861/1. Positie: 47° 56’ 56’’ N, 16° 1’ 15’’ E, 16° 1’ 15’’ E, 47° 56’ 56’’ N, Positie: 1861/1. grottenregistratienr. 14m verschil in altitude: 90m, de grot: van lengte 585m, altitude: deningeroides. 1. Druipsteengrot van Flatz, 2. Repolust grot, 3. Herkova 3. Repolust grot, 2. Druipsteengrot Flatz, van 1. deningeroides. Langes Loch) Oostenrijks (syn.: Tropfsteinhöhle Flatzer Gegevens: jama. Bergfreunde“). There is no informationThere sequence. about the layer Bergfreunde“). Kaart Oostenrijk Ursus van vindplaatsen van met Figuur 1: Geology: Steinalm limestone (Middle Triassic) Steinalm limestone (Middle Geology: of people („Die a local group 1904 to 1906 by Excavations: Position: 47° 56’ 56’’ N, 16° 1’ 15’’ E, altitude: 585m, length of cave: length of cave: 585m, altitude: 16° 1’ 15’’ E, N, 47° 56’ 56’’ Position: 14m difference: altitude 90m, register nr. 1861/1 nr. register 1 Dripstone cave of Flatz, 2 Repolust cave, 3 Herkova jama 2 Repolust cave, 1 Dripstone cave of Flatz, Austrian cave Langes Loch) (syn.: Tropfsteinhöhle Flatzer Basics: Figure 1: Map of Austria of Ursus with sites deningeroides of Map Figure 1:

] 8 [ ICBS PROCEEDINGS incisura media, inc. mes: incisura mesialis, me: mesialedge, mes: Abbreviations: de: distaledge, did: distoconid, dist: distal, inc.med: 5: i2sinFl-N17U. ingressusRabederetal. 2004(morphotype“s”) 4: i2sinFl-N32U. deningeroidesMottl, 1964(morphotype“d/s”) 3: i2sinFl-N34U. deningeroidesMottl, 1964(morphotype“d”) morphotype “d”, 2: morphotype“s” 1-2: schematicdrawings ofthelingualsite(afterRabeder, 1999). 1: cave, oftheFlatzer dripstone bears lingualview, 1,5x enlargement Figure 2: (i2)fromMiddleandLatePleistocene Secondlower incisors [ and theM2sup. occlusal pattern of theteethinparticulari2 inf.,i3inf. Flatzer caveisbased onthedimensionsand primitive SYSTEMATIC POSITION the smallerM2inmaxillaryfragment (F1-2). sup. of M2 sup.ofboth logically primeval.Fig.4.showsthedifferences betweenthe mesialis mesiaal, msd: mesioconid, prd: protoconid, sd: sulcusdistalis, sm: sulcus incisura media, inc. mes: incisura mesialis, me: mesiale zijde, mes: Afkortingen: de: distalezijde, did: distoconid, dist: distaal, inc.med: 5: i2sinFl-N17U. ingressusRabederetal. 2004(morphotype“s”) 4: i2sinFl-N32U. deningeroidesMottl, 1964(morphotype“d/s”) 3: i2sinFl-N34U. deningeroidesMottl, 1964(morphotype“d”) 1999). 1: Morphotype“d”, 2: morphotype“s” 1-2: schematischetekeningen van delingualezijde(naarRabeder, 1,5x vergroting van Flatz.pleistocene berenuitdedruipsteengrot Linguaalaanzicht, Figuur 2: Tweede (i2)van midden-enlaat- onderkaakssnijtanden mesialis mesial, msd: mesioconid, prd: protoconid, sd: sulcusdistalis, sm: sulcus 10 The taxonomicpositionofthesmaller bearteethfromthe The rarerecordedmolarsarerelativelysmallandmorpho Molars ]

Ursus cf. ingressus Ursus- species particularlyclear. The M2 has significantlymorecuspsthan

DORIS NAGEL - contemporaneous: and blackcrust),followingtaxacanbeconsideredto ASSOCIATED FAUNA sm: sulcusmesialis incisura media, me: mesialezijde, prd: protoconid, sl: sulcuslateralis, Afkortingen: de: distalezijde, did: distoconus, dist: distaal, inc.m: 3: i3dex Fl-N29, cf. Ursus ingressus 2: i3sinFl-N29, deningeroides Ursus 1: i3dex Fl-N29, deningeroides Ursus Stadtmuseum Neunkirchen 3-5: van Flatz, uitdedruipsteengrot i3infvan Ursus drie collectievan 1999). 1: morphotype A (sin), 2: morphotypeD(sin). 1-2: schematischetekeningen van delingualezijde(naarRabeder, 1,5x vergroting Figuur 3: (i3)van Ursus, Derdeonderkaakssnijtand linguaalaanzicht, mesialis media, me: mesialedge, prd: protoconid, sl: sulcuslateralis, sm: sulcus Abbreviations: de: distaledge, did: distoconid, dist: distal, inc.m: incisura 3: i3dex Fl-N29, cf. Ursus ingressus 2: i3sinFl-N29, deningeroides Ursus 1: i3dex Fl-N29, deningeroides Ursus Stadtmuseum Neunkirchen 3-5: caveofFlatz, fromDripstone threei3infofUrsus collection of morphotype A (sin), 2morphotypeD(sin) 1-2: schematicdrawings oflingualside(afterRabeder, 1999). 1 1,5x Figure 3: Third lower incisor(i3)ofUrsus, lingualview, enlargement • • • • • • • • Due tothetypicalconservationstatus(yellowwithrusset Hystrix vinogradovi ?Bison/Bos gnawing marksof Hystrix) Cervus elaphus Dama Sus scrofa L. Rhinocerotide indet.(fragmentof molar) 11 cheekteeth,1coffinbonewith gnawingmarks) Equus Ursus deningeroides sp. cf. mosbachensis

(1 fragmentofantler, molars) sp. (2 incisors) L.(fragmentsofantlerspartlywith Argyropulo, 1941(1mandible Mottl1964 Reichenau, 1903(1incisor,

R. GOLD CRANIUM

JUNI 2016 ICBS PROCEEDINGS

JUNI 2016 - GOLD R.

- - - - - Lem Lagopus,

, small Ursus in Ursus denin (= ?small carnivors (Panthera leo CRANIUM pelaeus“ ) remains of marmots are ) remains of marmots Marmota marmota

cf. s probably migrated to Middle cave bears (likely Bovidae indet. The chronological affiliation of ) positions them into the Middle :“Ursus Panthera leo spelaea), U. ingressus U. ingressus does not belong to this fauna association but is does not belong to this fauna association but Martes. Meles, Vulpes Cervus elaphus, , etc. that are assigned to the Late Glacial (about etc. that are assigned to the Late Glacial (about ), cave lion ( Spalax The conservation status of the small carnivores remains The conservation status of the small carnivores There is no radiometric dating but Thenius (1949) as Thenius (1949) radiometric dating but There is no and domestic and wild birds Holocene: bears, cave lion Late Pleistocene: cave Middle Pleistocene? we are of the opinion that the Due to the existing material Late Glacial period (OIS2): (OIS3): Middle Wurmian

Martes. Meles, Vulpes). Martes. Meles, Vulpes). Martes, Meles, Vulpes

gressus ( the same con cave bear and cave lion remains arises from servation status. extinct 29.000 Europe about 50.000 years ago and became ago. ( as well. Wurmian sumed that the vertebrate remains of the Flatzer dripstone of the Flatzer dripstone the vertebrate remains sumed that periods: to three different cave belong Bison, Capreolus, Cervus, Vulpes, spelaea), Martes, Meles, Bos, Marmota, Lepus geroides) stages originate from three different fossil vertebrate remains of the Pleistocene: carnivores ( Alpine caves but they are not from the cave often found in period most bear period but from the much younger cold Alpine caves There are plenty of probable the Late Glacial. with some (including the Merkenstein cave close to Flatz) like times massive presence of boreo-Alpine elements nivalis, Lepus timidus, Microtus mus, Dicrostonyx, Marmota & Rabeder 20-12ka BP) by radiometric dating (Döppes 1997). element. an interglacial CHRONOLOGY R. GOLD R. Martes, Panthera leo , (1 mandible)

ingressus Bos. cf. and Nordmann, 1840 Ursus may be definitely younger (Late Pleistocene). may be definitely younger (Late Pleistocene). The remains of spelaea Not clearly assignable are the remains of Bison Meles, Vulpes, dex with p4-m3, 3 I sup, gnawing traces, see Fig.5) dex with p4-m3, 3 I sup, gnawing traces, see Spalax leucodon • • 2: Rechter onderkaak (Fl- N69) van het stekelvarken Hystrix vinogradovi Argyropulo, 19xx (1,5x vergroting). Argyropulo, Hystrix Rechter onderkaak het stekelvarken vinogradovi (Fl- N69) van 2: Cervus (Fl-xx) van op een geweifragment een stekelvarken elaphus (1,5x vergroting) knaagsporen van van foto Vergrote 3: Midden-pleistocene overblijfselen en sporen van stekelvarkens uit de druipsteengrot Flatz stekelvarkens van en sporen van Midden-pleistocene overblijfselen grootte. ware Cervus Fl-xx van Geweifragment stekelvarkens, elaphus met knaagsporen van 1: 2: mandible dex (Fl- N69) of the porcupine Hystrix vinogradovi Argyropulo, 19xx (1,5x) Argyropulo, of the porcupine Hystrix (Fl- N69) dex vinogradovi mandible 2: enlarged picture of gnawing marks (Fl-xx) of Cervus of porcupine on an antler fragment 3: elaphus (1,5x) Fig. 5. Middle Pleistocene remains and traces of porcupine from the dripstone Middle Pleistocene remains and traces cave of Flatz 5. Fig. Fl-xx of Cervus size antler fragment natural elaphus with gnawing marks1: of porcupine, 2004 (Laat-Pleistoceen) (Midden-Pleistoceen) ingressus Rabeder et al. Ursus cf. invers), M2 germ Fl-Nx (dex, 2: vergroting 1,5x 1964 Ursus Mottl, deningeroides (sin), Fl-2 M2 in bovenkaak 1: 2004 (Late Pleistocene) occlusaal aanzicht, M2 sup uit de druipsteengrot Flatz, van 4. Fig. 2: M2 germ Fl-Nx (dex, invers), Ursus cf. ingressus Rabeder et al. ingressus Rabeder et al. Ursus cf. invers), M2 germ Fl-Nx (dex, 2: 1: M2 in maxillary Fl-2 (sin), Ursus deningeroides Mottl, 1964 (Middle 1964 Ursus M2 in maxillary Mottl, deningeroides Fl-2 (sin), 1: Pleistocene) Fig. 4. M2 sup from the dripstone cave of Flatz, occlusal view, occlusal view, M2 sup from the dripstone cave of Flatz, 4. Fig. enlargement 1,5x

] 10 [ ICBS PROCEEDINGS [ van Flatz deningeroides uitdedruipsteengrot Tabel 2: Afmetingen enmorphotypevan snijtandenvan Ursus caveofFlatz deningeroides fromthedripstone Table 2: ofUrsus Measurementsandmorphotypesofincisors 12 NK NK NK NK NK NK NK NK NK NK NK NK Fl-N46 Fl-N45 Fl-N45 FlN9+14 NK NK NK NK NK NK NK ] coll. coll. inv. nr. N128 Fl- Fl-N9 Fl-N29 Fl-N12 Fl-N11 Fl-N34 Fl-N32 Fl-N31 Fl-N33 Fl-N28 Fl-N39 Fl-N36 Fl-N38 Fl-N37 Fl-N5 N45 Fl- N22 Fl- N13 Fl- Fl-45

inv. nr. inv. nr. mandible mandible mandible mandible caninus Cinf-md. caninus caninus Cinf-md. sup I3 sup I3 i3 inf i3 inf i3 inf i2 inf i2 inf i2 inf sup I1,2 sup I1,2 i1 inf i1 inf i1 inf i1 inf element element element sin sin dex sin sin sin sin sin dex dex sin sin dex sin dex sin sin dex 20,7 21,9 23,6 ca. 24,4 22,4 side length of side crown 16,2 17,4 11,5 11,5 11,3 8,5 8,4 8,9 7,8 9,5 5,6 5,4 5,3 5,1 fr fr fr length 298 14,5 16,2 16,0 18,3 15,4 width of condylar worn 13,1 10,2 10,7 10,4 10,6 9,1 9,7 10,7 9,8 7,7 7,5 7,5 7,3 crown length width 21,8 - 24,8 27,8 - length of 178 167 fr fr ≥ >30,0 28,4 26,5 27,3 25,7 worn 46,1 36,4 ≥ 35,3 fr 29,8 ≥ 25,7 root 32,0 31,0 C-m3

14,5 - 17,8 17,3 - height width of 90,3) (95,3 94,6) fr d0 d2 A A A B root 0 0 A B A d d/s worn p4-m3

♀ ♀ ♀ ♂ ♀ morpho- sex type (66,8 (72,2 70,8 fr p4-m2 deningeroides van de druipsteengrot van Flatz deningeroides van dedruipsteengrot Tabel 3: Afmetingen enmorphotypesvan hoektanden van Ursus caveofFlatz deningeroides fromthedripstone Table 3: Measurements andmorphotypesofcaninesUrsus in bovenkaak) van Flatz(md:deningeroides uitdedruipsteengrot inonderkaak; mx: Tabel 4: Afmetingen enmorphotypesvan kiezen van Ursus in maxilla) caveofFlatz(md:deningeroides fromthedripstone inmandible; mx: Table 4: Measurementsandmorphotypesofcheek teethofUrsus 75,3) 78,7 78,4 fr Fl-N45 Fl-N45 Fl-N18 Fl-2 Fl-N11 Fl-N45 Fl-N4 Fl-N45 Fl-N45 Fl-6 Fl-N19 Fl-2 N9+14 Fl- Fl-N45 Fl-N45 Fl-5 Fl-N3 Fl-2 Fl-4 Fl-N69 N9+14 Fl- Fl-N45 m1-3 inv. nr. (52,0 54,5 54,8 57,9 m1-md m1inf P4-mx P4sup p4-md p4inf m3-md m3inf m3inf M2-mx m2-md m2-md m2-md m2inf m2inf M1-mx M1sup M1sup m1-md m1-md m1-m2 element 64 52,8 ca. fr 37,8 dex dex dex sin sin dex dex dex dex sin sin sin dex dex dex sin dex dex sin sin height side below m1 27,1 27,5 18,5 17,7 15,4 15,7 24,2 24,8 24,5 39,1 29,6 28,2 27,9 26,1 25,5 25,4 25,2 24,5 28,3 26,6 druipsteengrot van Flatz druipsteengrot deningeroidesuitde van Ursus Tabel 1: Afmetingen van onderkaken caveofFlatz dripstone deningeroidesfromthe of Ursus Table 1: Measurementsofmandibles length 12,7 14,3 13,1 12,5 8,2 8,3 18,0 18,8 17,6 20,6 18,2 18,4 16,3 15,2 17,6 17,9 18,0 17,6 13,5 13,1 width 12,7 14,3 13,1 12,5 8,2 8,3 18,0 18,8 17,6 20,6 18,0 18,4 16,3 15,2 17,6 17,9 18,0 17,6 13,5 13,1

width - - - B1 B worn worn B worn - - - worn worn worn A/B A/B A B1 B1

morpho-type CRANIUM

JUNI 2016 ICBS PROCEEDINGS - JUNI 2016 -

- - - - - . Mit - Docu

Annalen Quater , 161-165. 10 Internation

th group) to 19 9ème Sympo Mitteilungen der Cahiers scientific / The problem of

Early Pleistocene bear CRANIUM 49-67. , , 305-327. ° 2 Sitzungsberichte / Öster n (2013) , 1-301.

59-70. Ursus spelaeus (2011) (2011) , 1-102 , 1–56. 59, 1-4,

26

, 287-290. , 241-284. Pleistocene vertebrate faunas of Hungary Die Höhle 158, 4

Bärenphylogenese in Südost-Österreich. , 1-135. , 15-25. Mitteilungen der Kommission für Quartärfor Mitteilungen der Kommission 24/2

70. 17

deningeroides from the dripstonedeningeroides cave of Flatz een metatarsale UrsusAfmetingen van van 5: Tabel uit de druipsteengrotdeningeroides Flatz van Table 5: Measurements of a metatarsal Measurements Ursus bone of 5: Table

Geologija (La fauna pléistocene de la cavité de Jama pod Herkovimi (La fauna pléistocene de la cavité de Jama pod Herkovimi dd Pohar, V., I. Debeljak, G. Rabeder (2003) Cave bear site Jama pod I. Debeljak, V., Pohar, G. (1999) Die Evolution des Höhlenbärengebisses. Rabeder, (2004) New Withalm D. Nagel, G. G., M. Hofreiter, Rabeder, (2008) Withalm G. G., I. Debeljak, M. Hofreiter, Rabeder, Withalm G. G., M. Pacher, Rabeder, Thenius, E. (1949) Der erste Hinweis einer fossilen Blindmaus Argant, A. (1991) Carnivores Quaternaires de Bourgogne. Carnivores Quaternaires de Bourgogne. A. (1991) Argant, C. A. Gallay, H. Floss, Argant, A. Barriquand, L., L. Barriquand, (eds.) (1997) Pliozäne und pleistozäne Döppes, D., G. Rabeder G. Rabeder Alscher, Frischauf, C., M. D. (1986) Janossy, Mottl M. (1964) Mottl, M. (1967) Neues vom Hystrix-Horizont Europas. favna iz Jama pod Herkovimi (1981) Pleistocenska V. Pohar, Cave Bear Symposium Semriach (Styria, Austria). Abstracts & Austria). Cave Bear Symposium Semriach (Styria, 13,8

Herkovina pecmi (N. Slovenia): preliminary study. Herkovina pecmi (N. Slovenia): preliminary study. (Savoie, sium International Ours des Cavernes, Entremont-le-Vieux France), Abstr., Österreichischen teilungen der Kommission für Quartärforschung, 11 Akademie der Wissenschaften Alpine Cave Bears (Ursidae, ). of Taxa Hors série Dép. Rhône - Museum Lyon, Morphological response of cave bears ( high-alpine habitats. Austria). (Lower remains from Deutsch-Altenburg Akademie der Kommission für Quartärforschung, Österreichischen Wissenschaften in Österreich. (Spalax hungaricus Nehr.) Mathematisch-Naturwis Akademie der Wissenschaften, reichische und Abteilung I, Biologische Wissenschaften senschaftliche Klasse Erdwissenschaften, REFERENCES de géologie Lyon ments des laboratoires Qunif (2011) Y. C. Nykiel, Guérin, L. Guillot, M. Jeannet, 4 Hors-série naire Katalog der wichtigsten Fossilfundstellen Faunen Österreichs. Ein und ihrer Faunen. Akademie der Wissenschaften schung, Österreichischen Azé 1 and Herkova jama. Ursus deningeroides from al Excursions, 13-14. Akadémiai Kiadó, Budapest. am Geologie und Technik Mitteilung des Museums für Bergbau, Landesmuseum “Joanneum” 71 Museums in Wien des Naturhistorischen pecmi. pecni). dw 15,3 - - - : -

-

- U. et dew

Ursus

17,4 sdd

8,4

swd U. deningeri sp. are also ele

11,5 pd Dama 22,5 Hystrix vinogradovi,

and pw , a big horse, a small primeval 11,8 mosbachensis, Cervus elaphus, mosbachensis, Cervus

cf.

. 2013). The material of the Flatzer The . 2013). length As typi similar to the Flatzer fauna. Mottl differs from Mottl differs Hystrix 59,4

et al side Spalax- dex from the Repolust cave and for a contempora also belongs to this fauna association. Based also belongs to this fauna association. Based

like bear as well as interglacial elements such as elements as interglacial as well bear like

. element and

mt2 inv. nr. inv. The main part of the studies was made in the Institute of The study of the fauna was funded by the Lower Austria The study of the fauna was funded by the Lower For the possibility to study the fossils of the Flatzer Porcupine remains from Middle and Late Pleistocene Porcupine remains from Middle and Late Pleistocene Ursus deningeroides The similarities to the Repolust-fauna indicates that The similarities to the Repolust-fauna indicates Spalax leucodon, Sus scrofa The summary of this community primarily bases on the The summary of this community Warm period of Middle Pleistocene (OIS7 or OIS5?) (OIS7 or period of Middle Pleistocene Warm . 2011, Frischauf . 2011, Fl-15 are grateful to Mark and Elisabeth Procter for correcting the English. are made by Mr. Rudolf Gold (Institute of Palaeontology, Rudolf Gold (Institute of Palaeontology, are made by Mr. We and therefore we want to thank him too. Vienna) Univ. Palaeontology of the University of Vienna which especially Vienna Palaeontology of the University of Doris Nagel made possible. Some of the pictures Prof. Dr. government within the project “Lunzer Bärenhöhlen” for Erwin Pröll. which we would like to thank governor Dr. as Mr Benedikt Wallner from the Municipality Museum of from Wallner as Mr Benedikt Neunkirchen. dripstone cave we would like to thank Dr. Erich Steiner dripstone cave we would like to thank Dr. Austrian State Museum of St. Pölten as well from the Lower the Late Middle Pleistocene are possible too. ACKNOWLEDGMENTS not clear whether the “Hystrix-faunas” of Hungary, Austria not clear whether the “Hystrix-faunas” of Hungary, of and Slovenia are contemporaneous because interglacials Hystrix-horizon “Varbó”-phase and associate it chronologi Hystrix-horizon “Varbó”-phase cally into the beginning of the Late Pleistocene. Indeed it is shelter and Poroslyuk (Hungary). Janossy (1986) named the cal faunas of the Hystrix-horizon Mottl (1967) and Janossy Tarkö-rock (1986) instance the Lambrecht-Kálmán cave, “deningeri”- Sus scrofa horizon” (Mottl, 1967). This is characterized by the simul This horizon” (Mottl, 1967). taneous appearance of sediments of Middle Europe are very conspicuous because sediments of Middle Europe are very conspicuous name “Hystrix of their recent distribution which led to the deningeroides deningeroides the associated fauna. by neous existence is offered dripstone cave is too rare to determine the evolutionary level. dripstone cave is too rare to determine the evolutionary those bears are closely related to that strong argument A the higher evolutionary level of the premolars (Rabeder the higher evolutionary level of the premolars al from Reichenau in the clearly smaller dimensions and in from Reichenau in the clearly smaller dimensions horses a Middle Pleistocene age is to assume. deningeroides deningeroides but also of the on the primeval characteristics of those bears ments with an interglacial character. ments with an interglacial the Pleistocene, as indicators for a warm climate. the Pleistocene, as indicators tropical distribution and therefore they are classified, also for tropical distribution and the surroundings of the cave simultaneous with the Cervidae the surroundings of the nowadays have a tropical to sub and Equidae. Porcupines Repolust cave (Mottl, 1967, Döppes & Rabeder 1997). The 1967, Döppes & Rabeder 1997). Repolust cave (Mottl, porcupines indicate that they inhabited gnawing marks of the great similarity of the Middle Pleistocene fauna from the great similarity of the Dama dama Ursus deningeroides, Sus scrofa, Sus scrofa, Ursus deningeroides, Spalax leucodon, Equus

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