Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears


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Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears

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Title:
Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears
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Bulletin of Geosciences
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Diedrich, Cajus G.
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English

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Subjects / Keywords:
Panthera Leo Spelaea (Goldfuss, 1810) ( local )
Steppe Lion Skeletons ( local )
Late Pleistocene ( local )
Czech Republic ( local )
Taphonomy And Pathology ( local )
Palaeobiology ( local )
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serial ( sobekcm )

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Abstract:
The world’s first mounted "skeletons" of the Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) from the Sloup Cave hyena and cave bear den in the Moravian Karst (Czech Republic, central Europe) are compilations that have used bones from several different individuals. These skeletons are described and compared with the most complete known skeleton in Europe from a single individual, a lioness skeleton from the hyena den site at the Srbsko Chlum-Komín Cave in the Bohemian Karst (Czech Republic). Pathological features such as rib fractures and brain-case damage in these specimens, and also in other skulls from the Zoolithen Cave (Germany) that were used for comparison, are indicative of intraspecific fights, fights with Ice Age spotted hyenas, and possibly also of fights with cave bears. In contrast, other skulls from the Perick and Zoolithen caves in Germany and the Ur?ilor Cave in Romania exhibit post mortem damage in the form of bites and fractures probably caused either by hyena scavenging or by lion cannibalism. In the Srbsko Chlum-Komín Cave a young and brain-damaged lioness appears to have died (or possibly been killed by hyenas) within the hyena prey-storage den. In the cave bear dominated bone-rich Sloup and Zoolithen caves of central Europe it appears that lions may have actively hunted cave bears, mainly during their hibernation. Bears may have occasionally injured or even killed predating lions, but in contrast to hyenas, the bears were herbivorous and so did not feed on the lion carcasses. The articulated lion skeletons found in cave bear dens deep within caves scattered across Europe (such as those from the Sloup, Zoolithen and Ur?ilor caves) can therefore now be explained as being the result of lions being killed during predation on cave bears, either by the cave bears defending themselves or as a result of interspecific fights.
Original Version:
Bulletin of Geosciences, Vol. 86, no. 4 (7/3/05).

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!" #$# #%& ' Theworld’sfirstmounted“skeletons”oftheLatePleistocene Pantheraleospelaea (Goldfuss,1810)fromtheSloup CavehyenaandcavebeardenintheMoravianKarst(CzechRepublic,centralEurope)arecompilationsthathaveused bonesfromseveraldifferentindividuals.Theseskeletonsaredescribedandcomparedwiththemostcompleteknown skeletoninEuropefromasingleindividual,alionessskeletonfromthehyenadensiteattheSrbskoChlum-KomínCave intheBohemianKarst(CzechRepublic).Pathologicalfeaturessuchasribfracturesandbrain-casedamageinthese specimens,andalsoinotherskullsfromtheZoolithenCave(Germany)thatwereusedforcomparison,areindicativeof intraspecificfights,fightswithIceAgespottedhyenas,andpossiblyalsooffightswithcavebears.Incontrast,other skullsfromthePerickandZoolithencavesinGermanyandtheUrilorCaveinRomaniaexhibitpostmortemdamagein theformofbitesandfracturesprobablycausedeitherbyhyenascavengingorbylioncannibalism.IntheSrbsko Chlum-KomínCaveayoungandbrain-damagedlionessappearstohavedied(orpossiblybeenkilledbyhyenas)within thehyenaprey-storageden.Inthecavebeardominatedbone-richSloupandZoolithencavesofcentralEuropeitappears thatlionsmayhaveactivelyhuntedcavebears,mainlyduringtheirhibernation.Bearsmayhaveoccasionallyinjuredor evenkilledpredatinglions,butincontrasttohyenas,thebearswereherbivorousandsodidnotfeedonthelioncarcasses.ThearticulatedlionskeletonsfoundincavebeardensdeepwithincavesscatteredacrossEurope(suchasthose fromtheSloup,ZoolithenandUrilorcaves)canthereforenowbeexplainedasbeingtheresultoflionsbeingkilledduringpredationoncavebears,eitherbythecavebearsdefendingthemselvesorasaresultofinterspecificfights.€Key words: Pantheraleospelaea (Goldfuss,1810),steppelionskeletons,LatePleistocene,CzechRepublic,taphonomyand pathology, palaeobiology. DIEDRICH,C.G.2011.LatePleistocene Pantheraleospelaea (Goldfuss,1810)skeletonsfromtheCzechRepublic(cen tralEurope);theirpathologicalcranialfeaturesandinjuriesresultingfromintraspecificfights,conflictswithhyenas,and attacksoncavebears. BulletinofGeosciences86(4) ,817…840(13figures,2tables).CzechGeologicalSurvey,Prague. ISSN1214-1119.ManuscriptreceivedMarch15,2011;acceptedinrevisedformAugust29,2011;publishedonlineNo vember 9, 2011; issued November 16, 2011. CajusG.Diedrich,PaleoLogicResearchInstitute,Nansenstrasse8,D-33790,Halle/Westphalia,Germany;cdiedri@gmx.netTheLatePleistoceneEurasiansteppelionspecies,aclose relativetoanextinctsubspeciesofthemodernAfrican lion(fromDNA…seeBurger etal. 2004),wasfirstdescri bedandillustratedfromtheZoolithenCavesiteinBava ria,southernGermany,andclassifiedasthe“cavelion” Felisspelaea Goldfuss(1810).Thelargemaleholotype skull,withitspathologicalbitemarkfeatures,hasre centlybeenrelocatedandreclassifiedasthe“steppelion” (popularnamebyDiedrich2008) Pantheraleospelaea (Goldfuss,1810)(DNAbyBurger etal .2004).The steppelionpopulationfromthiscaveisthelargestknown inEurope;ithasrecentlybeenreviewedanddiscussedin abroadtaphonomicandpalaeobiologicalcontextrelating topredationoncavebearsandconflictswithhyenas(Die drich2010d,2011a). Followingthisdiscovery200yearsagootherimportant Pleistocenelionfindshavebecomefamoussuchastheold estmounted“cavelionskeletons”describedhereinfrom theSloupCave(Fig.1A,B)intheMoravianKarst(Czech Republic,centralEurope),ahyenaandcavebeardensite (Diedrich2009c,2011b).Thefirstofthetwo“skeletons” ofSloupCave( cf .Wankel1858;“skeleton2”…Brno, herein;Fig.1B)isexhibitedintheAnthroposMuseum, Brno.Wankel(1888)laterreferredtoasecondskeleton !

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(“skeleton1”…Vienna,herein;Fig.1A)andalsoillustrated anadditionalskullfromthesamecave.Thetwomounted “skeletons”ofthecavelion Felisspelaea werecompiled frommaterialfoundintheCut-StoneGalleryoftheSloup Cave( cf .Wankel1858,1868,1888;Fig.2).Thismaterial iscriticallyanalysedherein,togetherwithnewdiscoveries fromthe1997SloupCaveexcavations( cf .Seitl1998)that compriseanotherlargemaleskullandsomepostcranial bones(Diedrich2011).Wankel(1892)carriedoutfurther excavationsattheSloupCaveandreportedtwomore“lion skeletons”,onewithapathologicalconditiononthelower jaw,andtheotherdescribedasbeingalso“quitecom plete”,butwhetherthesestillexistandifso,wheretheyare stored, remains unclear. Lionremainshavealsobeenreportedfromthe MoravianKarstinotherpublications,particularlyfromthe SloupandVýpustekcavesbutalsofromothercaves,and arelistedbyMusil(1956).Apreliminaryoverviewofthe BohemianandMoravianKarstlionlocalitiesispresented herein(Fig.2).Theanalysisoflionbonematerialfrom thesesitesisincompleteandwarrantsfurtherinvestiga tions. LionremainshavebeenreportedfromtheTurská MatalCave(Kafka1903)andtheChlupáovaslujCave ontheKobylaHillnearKonprusy(Zázvorka1954),inthe BohemianKarstintheCzechRepublicconsistingofjust singlebonesandonesinglecubskeletonremainhavebeen published,fromwhichthesesiteshaverecentlybeeninterpretedtobebothhyenapreydepotsandhyenadens (Diedrich&™ák2006).Thematerialrequiresfurther study,however,asitincludesalioncubskeletonfromthe ChlupáovaslujCave.Themostcompleteknownlion skeletoninEuropefromasingleindividual(fromthe SrbskoChlum-KomínCaveintheBohemianKarst)had beenpreviouslyillustrated(Diedrich&™ák2006)butwas relocatedandstudiedindetailabout35yearslaterherein. Thissiteyieldedapproximately1,500boneremainsfrom horses,apparentlyaccumulatedbyahyenaclanthathad developedaspecializationinhuntinghorsesinresponseto thescarcityofmammothsinthemountainousregions(re strictedtotheBerounkaRivervalley)oftheBohemian Karst (Diedrich 2010a). LionremainsfromLatePleistoceneopenairsiteshave beenreportedfromlocalitiesaroundPrague(Vltava Riverterrace;Diedrich2007a)andBeroun(Berounka Riverterrace;Kafka1903,Diedrich2007a;Fig.3)inthe CzechRepublic.Thismaterial,whichincludesaskull, teeth,andisolatedskeletalbones,hasrecentlybeenre-de scribedtogetherwithmaterialfromseveralotheropenair localitiesinthenorth-westernCzechRepublic,including Trmice (Diedrich 2007a). SkeletonsoftheLatePleistocenesteppelion Panthera leospelaea havealsobeenreportedfromafewothersites inEurope.Theoldestknownskeletonisfromthecave beardenintheAzéCave(France),andhasbeendated intotheSaalianofthelateMiddlePleistocene(Argant 1988).Theskeletonofasenilelionesswithpathological features,whichwasfoundinanelephantgraveyardatthe GermanNeumark-Nord-Lake1site,wasfromtheEem ianinterglacialoftheearlyLatePleistocene(Diedrich 2010b,c).Thenumberofrecordedskeletonsthenin creasesduringthecoldperiodoftheWeichsel/Würmgla ciation(LatePleistocene).Alargelyintactskeletonofa large,strongmalelionwasfoundintheArrikrutzCavein Spain(Altuna1981),anotherwasdescribedfrom SalzburginAustria(Tichy1985),andathirdmaleskele tonwasfoundinanopenairsiteatSiegsdorfinGermany (Gross1992).Therecentdiscoveriesoflionskeletons deepwithintheRomanianUrilorCave(Diedrich etal. 2009)arethemostspectacularbecausetheyrepresentthe firstclearevidencethatsteppelionshuntedcavebears deepwithintheircaves(Diedrich2011f).SkeletalremainsfromtheZoolithenCavehavealsobeenreviewed recentlybutcompleteskeletonscouldnotbecompiled (Diedrich2010d). Themainobjectiveofthiscontributionistocritically re-examinethehistoricalSloupCavelionskeletoncompilations,togetherwiththebonesoftheonlyknownlioness skeletonfromtheCzechRepublic(fromtheSrbsko Chlum-KomínCave),whichisillustratedinabonecata logue(Figs6…9)forosteologicalcomparisons.Asecond objective,includingthishistorical(largelyunstudied)bone material,wastoexaminethebitedamageandotherpatho logicalfeaturesonthelionmaterialfromtheCzechRepub lic,withrespecttointerspecificandintraspecificantago nismandresultingskulldamage.Theseimportantsteppe lioncavefindsarecompared,mainlywithrespecttotheir taphonomyandpathologicalfeatures,tosteppelionmate rialfromothercaves(Diedrich2009a,b,20110d,2011e) andopenairsites,bothinGermanyandintheCzechRe public(Diedrich2007a,2010b,2011d;Diedrich& Rathgeber 2011). (#) TwofirstcompleteEuropean(composite)skeletonsoftheLatePleistocenesteppelion Pantheraleospelaea (Goldfuss,1810),whosebones weremainlyfoundintheCut-StoneGalleryoftheSloupCave,intheMoravianKarstoftheCzechRepublic(compositephalangesalsofromVýpustek Cave). €A…skeleton1( cf .alsoFigs2,3),whichwasdonatedin1885bythePrinceofLiechtensteintotheNaturalHistoryMuseuminVienna,Austria (NHMVNo.1885/0014/4302),inlateralview .€B… skeleton2( cf .alsoFigs2,3)intheAnthroposMuseumBrno,CzechRepublic(AMBwithoutNo.), whichhasseveralflaws(thepesaretransposed,theatlasistransposed,therearenosternalbones),inlateralandcranialviews(historicaldrawin gsfrom Wankel 1888).

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,&Thispaperexaminesthreelion“skeletons”fromtheSloup (MoravianKarst)andSrbskoChlum-Komín(Bohemian Karst)caves,togetherwithisolatedmaterialfromvarious caveandopenairsitesintheCzechRepublic.Dimensions andbitedamageonskullsarecomparedtosteppelion, hyena(hyenaprey),andcavebearmaterialfromother (#-) LatePleistocenesteppelionopenairandcavelocalitiesandhyenadensitesintheCzechRepublic,withdetailsfromCentralBohemia,Czech Republic (compiled from Diedrich & ™ák 2006; Diedrich 2007c, 2011, and the Sloup Cave map after Zajíek 2007).

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EuropeansitessuchasthePerickandZoolithencavesin GermanandtheUrilorCaveinRomanian.Historically collectedsteppelionmaterialfromtheSloupCaveishoused intheNaturalHistoryMuseum,Vienna(NHMV)compri singasingleskeleton(“skeleton1”…Vienna,herein)and someisolatedbones.TheSloupCavemegafaunaofthe WankelcollectionswasreceivedbytheNaturalHistory MuseumofViennain1885,formingoneofitsfirstpalae ontologicalcollections,buthadbeenexcavatedin 1881…1882forJ.WankelbyV.Sedlák(anemployeeofthe CzechProfessorK.Absolon).Thismaterialseemstohave beenusedmainlyfortheskeletonreconstructiondonated tothemuseumin1885bythePrinceofLiechtenstein.A thirdlionskullwasfromtheexcavationsbySeitlin1998, whichwasagainfoundintheCut-StoneGallerywherethe otherlionboneshadbeenfoundhistorically.Thismaterial isstoredintheAnthroposMuseum,Brno(AMB),asisthe second“skeleton”fromtheSloupCave(“skeleton2”… Brno,herein).TheSloupCave(“Slouper-Höhle”inGer man)andtheVýpustekCave(“KiriteinHöhle”inGerman) wereexploredbytheauthorpriortothe13thInternational CaveBearSymposiumin2007toidentifytheareasexcavatedhistoricallybyWankelandSedlak,andthemorerecent excavations by Seitl. Theunsystematicpickaxe-and-shoveldigsofthe SpeleoclubPrahain1968…1969causedextensivedamage toalionskeletonfromtheSrbskoChlum-KomínCave (“skeleton3”herein),nowhousedintheNaturalHistory Museum,Prague(NMP).Theskeletonwasfinallycompletedin2005withtheinclusionofmanybonefragments found36yearsafterthe“cavecleaning”,includingonerib fragment,threescapulafragmentsfrombothrightandleft scapulas,andoneupperjawM1tooth.Itisnowoneofvery fewalmostcompleteEuropeanLatePleistocenesteppe lionskeletonsfromasingleindividualandincludes149 bones(Table1).Thisskeletalmaterialwasseparatedfrom thatfromasecond,smallerskeletonofaoneyearoldcub, whichisrepresentedbyonly1/3ofitsbonesandexhibits typicallynon-fusedjoints.TheSrbskoChlum-KomínCave hasyieldedmorethen3,500bones,includingabout350 hyenaremains,aswellastheircoprolites,andalargequan tityofhyenaprey(dominatedbyhorseremains…including afoetalskeleton).Cavebearboneshavenotbeenrecorded in this Srbsko Chlum-Komín hyena den cave. Thedimensionsoflionskullsfromrecentlydiscovered caveandopenairsitesintheCzechRepublic,aswellas thosefromotherskeletalandskullremainsinEurope(in cludingthelongbonesdimensions),havebeencompared forasexidentificationpurposes.Bonesfromdistalextremi tiesarenotusefulforthispurposeastheyhavenotyetbeen studiedinsufficientquantities,similarasmetapodials. Finally,pathologicalfeaturesandbitemarkswereob servedonthehistoricalmaterialfromtheZoolithenCave inGermany,storedintheNaturalHistoryMuseumofthe HumboldtUniversity,Berlin(MB)andattheUniversityof Erlangen(UE),aswellasinmaterialfromtheSloupCave. Thepathologicalfeaturesandbitemarksindicate interspecificandintraspecificaggression,thusproviding importantcluesforunderstandingthepalaeoecologyof steppelionsandtheirrelationshipstohyenasandcave bears during the Late Pleistocene of Eurasia..&#&Order Carnivora Bowdich, 1821 Family FelidaeFischer, 1817 Subfamily Pantherinae Pocock, 1917 Genus Panthera Oken, 1816 Panthera leo spelaea (Goldfuss, 1810) Figures 1, 4…10, 12TheViennaskeletonisacompositeskeletonmadeupof bonesfromdifferentanimals.Itincludesbonesfrommale andfemaleanimalsaswellasseveralcastsofbones (Fig.3A).SomephalangesfromtheVýpustekCave(marked“Vý”or“Výpustek”; cf .Fig.4J,O)havealsobeenincludedinthepedalskeleton.Mostofthelargebonesand theskullofthisskeletonseemtobefrommales,possibly derivedfromanarticulatedmaleskeletonfoundinthe Cut-StoneGalleryoftheSloupCaveinhistorictimes, whilemanyoftheremainingbonesareeithercastsorfrom females. Skull.– Theskullisoriginalandfairlycomplete,withato tallengthof390mm( cf .Figs1A,4D,E),butitismissing severalteethsuchasbothP2,M1,andtheleftI3.Onlythe rootoftherightI2ispreserved,thetiphavingbeenflaked duringtheanimal’slifetime.TheP3lengthis28mmand theP4lengthis41mm.Theteetharehardlywornandthe skullsuturesarefusedinthe brain-casebutnot inthe nasals, provingthistohavebeenanadultindividual3…8yearsold. Abitemarkispresentonthesaggitalcrest(Fig.4E).The lowerjawsarepossiblyfromthesameskull;theleftman diblehasatotallengthof256mmandtheleftP4toothhasa length of 31 mm. Vertebralcolumnandpelvis.– Thefirsttwocervicalver tebrae(atlas,axes)areoriginal;theC3-6,andT1-2are casts.TheT3uptotheL4(lumbarvertebrae, cf .Fig.4L)are originals,suchastheL7,butallothervertebraearemoulds. Eightofthecaudalvertebraeareoriginalbones(Fig.4M),of whichthefirstseven arearticulated inanatomic correctrow !!"#$%&"'

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/ ) Bonesoftheadultfemale Pantheraleospelaea skeletonfromthehyenadenSrbskoChlum-Komíncavesite(CzechRepublic,BohemianKarst). No.Coll.-No.Bone typeCommentaryLeftRightCollection 1R 4406CraniumIncomplete, without both P2 and M1NMP 2R 4407/4408Lower jawsIncomplete, without I1xxNMP 3R 4415ScapulaIncompletexNMP 4Ra 4233ScapulaIncompletexNMP 5R 4417HumerusCompletexNMP 6R 4522HumerusCompletexNMP 7R 4418UlnaCompletexNMP 8R 4419UlnaCompletexNMP 9R 4420RadiusNearly completexNMP 10R 4421RadiusCompletexNMP 11R 4605ScapholunatumCompletexNMP 12R 4606ScapholunatumCompletexNMP 13R 4621PisiformNearly completexNMP 14R 4934PisiformNearly completexNMP 15R 4219ScaphoulnarCompletexNMP 16R 5150MetacarpusV, completexNMP 17R 5149MetacarpusIV, completexNMP 18R 5031MetacarpusI, completexNMP 19R 5148MetacarpusIII, completexNMP 20R 4535MetacarpusIV, completexNMP 21R 4536MetacarpusV, completexNMP 22R 4628PhalanxI, manus, digit VxNMP 23R 5029PhalanxI, manus, digit VxNMP 24R 4932PhalanxI, manus, digit IVxNMP 25R 4544PhalanxI, manus, digit IIIxNMP 26R 4920PhalanxI, manus, digit IIIxNMP 27R 4626PhalanxI, manus, digit IxNMP 28R 4935PhalanxII, manus, digit VxNMP 29R 4930PhalanxII, manus, digit IVxNMP 30R 4545PhalanxII, manus, digit IIIxNMP 31R 4927PhalanxII, manus, digit IIIxNMP 32R 4928PhalanxII, manus, digit IIxNMP 33R 4626PhalanxII, manus, digit IxNMP 34R 4933PhalanxIII, manus, ?digit IVxNMP 35R 4625PhalanxIII, manus, ?digit IIIxNMP 36R 5153PhalanxIII, manus, ?digit IVxNMP 37R 4926PhalanxIII, manus, ?digit VxNMP 38R 4523/4224/4565PelvicNearly complete, with sacrumNMP 39R 4525FemurNearly completexNMP 40R 4526FemurNearly completexNMP 41R 4602PatellaCompletexNMP 42R 4533PatellaCompletexNMP 43R 4528TibiaNearly completexNMP 44R 4527TibiaCompletexNMP 45R 4607FibulaWithout proximal jointxNMP 46R 4532FibulaHalf with distal jointxNMP 47R 4530CalcaneusCompletexNMP 48R 4914AstragalCompletexNMP 49R 4531AstragalCompletexNMP 50R 4937CuboidCompletexNMP

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/ ) continued No.Coll.-No.Bone typeCommentaryLeftRightCollection 51R 4635NavicularCompletexNMP 52Ra 4232Tarsal IIHalfxNMP 53R 4539MetatarsusV, completexNMP 54R 4538MetatarsusIV, completexNMP 55Ra 4231MetatarsusIII, proximal joint, fragmentxNMP 56R 4537MetatarsusII, completexNMP 57R 4543MetatarsusV, completexNMP 58R 4542MetatarsusIV, completexNMP 59R 4541MetatarsusIII, completexNMP 60R 4540MetatarsusII, completexNMP 61R 5038PhalanxI, pes, digit VxNMP 62R 4919PhalanxI, pes, digit IVxNMP 63R 4632PhalanxI, pes, digit IIIxNMP 64R 4627PhalanxI, pes, digit IIxNMP 65R 4922PhalanxI, pes, ?digitxNMP 66R 5030PhalanxII, pes, digit IIxNMP 67R 4942PhalanxII, pes, ?digitxNMP 68R 4929PhalanxII, pes, digit IVxNMP 69R 4633PhalanxII, pes, digit IIIxNMP 70R 5154PhalanxII, pes, ?digit IVxNMP 71R 4624PhalanxIII, pes, ?digit IIxNMP 72R 4923PhalanxIII, pes, ?digit IIIxNMP 73R 5151PhalanxIII, pes, ?digit IVxNMP 74R 4924PhalanxIII, pes, ?digit VxNMP 75R 5152PhalanxIII, pes, ?digit IIIxNMP 76R 4630PhalanxIII, pes, ?digit VxNMP 77R 4409Cervical vertebraAtlas, incompleteNMP 78R 4410Cervical vertebraAxis, incompleteNMP 79R 5032Cervical vertebraNo. 3, incompleteNMP 80R 4414Cervical vertebraNo. 4, incompleteNMP 81R 4411Cervical vertebraNo. 5, incompleteNMP 82R 4412Cervical vertebraNo. 6, incompleteNMP 83R 4413Cervical vertebraNo. 7, incompleteNMP 84R 4548Thoracic vertebraNo. 1, completeNMP 85R 4549Thoracic vertebraNo. 2, incompleteNMP 86R 4551Thoracic vertebraNo. 3, incompleteNMP 87Ra 4214Thoracic vertebraNo. ?, Proc. spinosusNMP 88R 4550Thoracic vertebraNo. ?, incompleteNMP 89R 4552Thoracic vertebraNo. ?, incompleteNMP 90R 4553Thoracic vertebraNo. 9, incompleteNMP 91R 4555Thoracic vertebraNo. 10, incompleteNMP 92R 4554Thoracic vertebraNo. 11, completeNMP 93R 4556Thoracic vertebraNo. 12, incompleteNMP 94R 4557Thoracic vertebraNo. 13, completeNMP 95R 4558Thoracic vertebraNo. 14, incompleteNMP 96R 4559Lumbar vertebraNo. 1, completeNMP 97Ra 4229Lumbar vertebraNo.2, two fragmentsNMP 98R 4561Lumbar vertebraNo. 3, incompleteNMP 99Ra 4230aLumbar vertebra?No. 4, fragmentNMP 100Ra 4230bLumbar vertebra?No. 4, fragmentNMP !!"#$%&"'

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/ ) continued No.Coll.-No.Bone typeCommentaryLeftRightCollection 101Ra 4230cLumbar vertebra?No. 5, 2 fragmentsNMP 102R 4563Lumbar vertebraNo. 6, incompleteNMP 103R 4564Lumbar vertebraNo. 7, incompleteNMP 104R 4566Caudal vertebraNo. ?, upper, incompleteNMP 105R 4567Caudal vertebraNo. ?, upper, completeNMP 106R4568Caudal vertebraNo. ?, upper, incompleteNMP 107R 4569Caudal vertebraNo. ?, upper, completeNMP 108R 4570Caudal vertebraNo. ?, middle, completeNMP 109R 4571Caudal vertebraNo. ?, middle, completeNMP 110R 4597Caudal vertebraNo. ?, middle, incompleteNMP 111R 4572Caudal vertebraNo. ?, middle, completeNMP 112R 4573Caudal vertebraNo. ?, lower, completeNMP 113R 4574Caudal vertebraNo. ?, lower, completeNMP 114R 5377Caudal vertebraNo. ?, lower, completeNMP 115R 5378Caudal vertebraNo. ?, lower, completeNMP 116R 5380Caudal vertebraNo. ?, lower, completeNMP 117Ra 4280Caudal vertebraLast two fusedNMP 118Ra 4224CostaNo. 1xNMP 119Ra 4225CostaNo. 2, completexNMP 120Ra 4226CostaNo. 3, completexNMP 121Ra 4227CostaNo. 4xNMP 122Ra 4228CostaNo. 5xNMP 123Ra 4248CostaNo. 6, completexNMP 124Ra 4230CostaNo. 7, completexNMP 125Ra 4232CostaNo. 8, completexNMP 126Ra 4233CostaNo. 9xNMP 127Ra 4234CostaNo. 10, completexNMP 128Ra 4235CostaNo. 11, nearly completexNMP 129Ra 4236CostaNo. 12, completexNMP 130Ra 4237CostaNo. 13, completexNMP 131Ra 4238CostaNo. 1xNMP 132Ra 4239CostaNo. 2xNMP 133Ra 4240CostaNo. 3xNMP 134Ra 4241CostaNo. 4xNMP 135Ra 4242CostaNo. 5xNMP 136Ra 4243CostaNo. 6xNMP 137Ra 4244CostaNo. 7xNMP 138Ra 4245CostaNo. 8xNMP 139Ra 4246CostaNo. 9xNMP 140Ra 4247CostaNo. 10xNMP 141Ra 4249CostaNo. 11xNMP 142Ra 4250CostaNo. 12xNMP 143Ra 4251CostaNo. 13xNMP 144Ra 4252Sternal boneNo. 1NMP 145Ra 4253Sternal boneNo. ?2NMP 146Ra 4220SesamoidCompleteNMP 147Ra 4221SesamoidCompleteNMP 148Ra 4222SesamoidCompleteNMP 149Ra 4223SesamoidCompleteNMP

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(possiblyfromoneindividual).The positionsofonecen tralandonedistalcaudalvertebraeareuncertain.Thecoxa is363mmlongandhasanacetabulumdiameterof50mm; thesacrumofthepelvisisamould.Therightribsinclude onefromacub(mostprobablyfromacavebear)witha non-fusedhead(the13thrib).Alloftheotherpreserved ribs, i.e .the5th,7th,9th(withpathologicalfeature,Fig.4K) 10th,11thand12th,havefusedribheads.Theleftribshave partlybeenpreserved(Nos.1,2,4,6,8,and11);othersare casts, as are all sternal bones. Appendicularskeleton.– Therightscapulaiscomplete (length312mm,glenoidwidth68mm,Fig.4F)buttheleft scapulaisacast.Bothlargehumeriaresimilarlyproportio nedandpreserved(length380mm,distalwidth100mm, Fig.4G,H).Neitherulnaisoriginalandonlythelargeright radiusiscomplete(withmaleproportions:length340mm, distalwidth75mm,Fig.4I).Allfiverightmetacarpiare originals(Fig.4J),whiletheleftmetacarpihaveacastof theMcI.Thedifferentproportionsandlackofsymmetry betweenleftandrightmetacarpalssuggestamixedderiva tionfrombothmaleandfemalespecimenswhich,insome cases,mayhaveevenderivedfromdifferentcaves( i.e .the SloupandVýpustekcaves; cf .Fig.4J).Alloriginalcarpaliaintherightmanusaremissingandonlyrepresentedby castswhileintheleftmanusthescapholunatae,andprobablyalsothemetacarpalbones,areoriginals.InbothforelimbmanusskeletonssomeofthephalangesI…IIIarecasts. Thehindlimbshavetwolargesymmetricalfemora(length 425mm,distalwidth90mm)andonelargerighttibia (length364mm,distalwidth68mm,Fig.4N).Bothoriginalcalcaneiaresymmetricalandlarge(128mminlength). Onceagain,themetatarsalsandphalangesdonotoriginate fromasingleindividualorasinglecavesite.Whereasmost bones are originals, the Mt I is a cast (Fig. 4O).Theskullofthisskeletonisacompilation,withatleastone mandiblederivedfromadifferentsource;thefrontlegsare lessthen50%madeupoforiginalbones,andthoseinthe hindlegsoriginatefrombothmalesandfemales.Itisun clearwhichpartsmighthavebelongedtotheoriginalske letonbecauseitissuchacompletemixture,includingma terialfromdifferentskeletons(includingbothmalesand females)fromtheCut-StoneGalleryoftheSloupCave, and there are even more casts than in skeleton 1 (Vienna). Skull .…Theoriginalskull(Fig.4A)hasatotallengthof 357mmandismissingsomeoftheoriginalteeth.Onthe rightandleftitlacksthesmallM1,whereastheleftsideca ninetoothisamuchtooshortcast.Thefrontalwidthis 98mm;theleftP4lengthis36mm.Thelowerjawappears tobeacompositefrommorethanoneoriginal.Theright mandible,withitscompletedentition,isconsistentwitha youngadultanimalskullinwhichtheanteriorpartsare non-fused,butthebrain-casebonesarefused.Theleft mandibleisfromanolder,senileindividual,withanM1that is highly worn, as are also the P3…4teeth. Vertebralcolumnandpelvis .…Theatlashasbeenmounted thewrongwayround;theaxisvertebraisanoriginal.All # (#0) Compositeand“real”skeletonsoftheLatePleistocenesteppe lion Pantheraleospelaea (Goldfuss,1810)fromCzechRepubliccave sites .€A… compositeskeleton1withbonesfromtheSloupCave,inthe NaturalHistoryMuseum,Vienna(NHMVNo.1885/0014/4302, cf . Fig.1A) .€B… compositeskeleton2withbonesfromtheSloupCave,on displayintheAnthroposMuseum,Brno(AMBwithoutNo.).€C…single skeletonfromtheSrbskoChlum-KomínCave,intheNaturalHistoryMu seum, Prague (NMP No. R 4406) (red = original bones, white = casts).* + !!"#$%&"'

PAGE 10

thecervicalvertebraearecasts.The3…7,9,11,13and14 (last)thoracicvertebraeareoriginals,asistheentirelum barvertebralcolumnL1-7whichmaythushavederived fromasingleindividual.Sixteenofthecaudalvertebrae areoriginalsbutthedistalvertebraearecasts.Noneofthe ribs are original; all of the sternal bones are missing. Appendicularskeleton .…Theleftscapulaisnotoriginal butafragment(includingtheglenoid)ispreservedfrom therightscapula.Thelefthumerusiscompleteand 362mmlong,whileonlythedistalhalfoftherighthume rusispreserved(distalwidth92mm).Theleftradiusis missingtheanteriorjointbuttherightradiusiscomplete (length324mm,Fig.4B).Bothulnaearecasts,asaremost ofthecarpals.Theleftmanushastheoriginalpisiform,and possiblyoneotheroriginalcarpalbone.Themetapodialsin themanusskeletonsarecompositesthatappeartoallbe fromtherighthandside,withtheMcII…Vhavingbeende rivedfromasingleindividual.AllphalangesI…IIareorigi nals,asarethreephalangesIII…thefourthisacast.The rightmanusagainhasoriginalMcII…Vmetapodials,asare allphalangesI…II,butonlythemiddletwophalangesIII arebones,theouterphalangesarecasts.Thehindlimbsare inbothcasesmadeupoforiginalfemoraandtibiaebones buttheyarenotsymmetricalanddonothavethesamepro portions,suggestingthattheydonotcomefromasinglein dividual.Theleftfemurislonger(440mm)thantheright femur(380mm).Incontrast,therighttibiaisonly345mm long,theleftoneis362mm,provingthesetobecomposi tesmadeupofbothmaleandfemalehindlimbbones.The ! (#1) Selectedbonesfromskeletons1 and2oftheLatePleistocenelion Panthera leospelaea (Goldfuss,1810),fromtheCutStoneGalleryoftheSloupCave,Moravian Karst(CzechRepublic).€A…skeleton2 (Brno)skull( cf .Fig.1A),lateral. €B…skele ton2(Brno)leftradius,lateral. €C…skeleton 2(Brno)pelviswithlargecarnivorebite marks( cf .Fig.3).€D…skeleton1(Vienna) skull,frontal .€E… skeleton1(Vienna)skull, detailofthepathologyontheleftsideofthe brain-case( cf .alsoFig.12B),lateralleft. €F…skeleton1(Vienna)rightscapula,lat eral .€G… skeleton1(Vienna)lefthumerus, lateral. €H…skeleton1(Vienna)righthu merus,lateral .€I… skeleton1(Vienna)right radius,lateral. €J…skeleton1(Vienna)right compositemanusskeleton,cranial .€K…skel eton1(Vienna)middleribwithpathology, lateral .€L… skeleton1(Vienna)middle lumbarvertebrae,lateral .€M… skeleton1 (Vienna)middlecaudalvertebrae,lateral. €N…skeleton1(Vienna)righttibia,cranial. €O…skeleton1(Vienna)compositeleftpes skeleton, cranial.* 2 ( ! 3 , 4 5 6 7 8 +

PAGE 11

feetareincorrectlymounted.Therightcalcaneushasproximalbitemarksandisincomplete,whiletheleftcalcaneusiscomplete.Theleftpesskeletonhasalltarsalia,the MtII…V,phalangesI…II,andthreephalangesIII;theinner phalanxIIIisacast.ThemissingdigitIhasnotevenbeen reproducedoneitherfoot.Therightpesskeletonconsists ofalltarsalia,theMtII…V,3phalanxIandallphalanxII; withinthephalanxIIIonedigitIIisacast.Allthephalan gesofthepedalskeletonsseemtobecompositesandoften phalanxIIisinthewrongposition,asaresomeoftheother phalanges. !Thisskeletonhas149bonesthatbelongtotheoriginalskele tonofayoungadultlioness(Fig.5,Table1),whereasthe phalangeIIIandsomephalangeIImay,insomecases,have derivedfromasecondskeletonofajuvenileanimal.The distalextremitybones(startingfromthemetapodials)are thereforenotillustratedherein.Allothernon-pedalbones havelargerdimensionsandbelongtotheoriginalskeleton illustratedhereinofayoungadultlioness.Noscavenging marksarepresentonthebones.Manymodernfracturesare presentandsomepiecesofbonearemissing,allofwhichare theresultofcarelessexcavation.Evidencethatthebonematerialisfromasingleindividualisprovidedbythesymmetry oftheskeleton,theconsistentproportions,andthelackof anyrepetitioninthebones.Theoriginofthesmallpedalbo nes(phalanges)ismoreproblematicandthesemayhave comefromasecond,juvenileskeleton. Skull .…Inmanyoftheviewsillustratedhereinthecranium withlowerjaw(Fig.6)canbeseentobeextensivelydama ged.BothupperjawP2aremissing,asistheleftM1.The incisors,bothcanines,andtheP3…4areintheiralveoli.The nasalbonesandfrontoftheskullhavebeendamaged.The lowerjaw(Fig.6F,G)hasbeenbadlydamagedbutboth mandiblesfittogetherexactlyintheirsymphyses.The rightmandibleincludestheI2…3,C,andP3…4;theleftman diblehasnoincisorsinitsalveoli.Bothmandiblesarein completeinthevicinityoftheramus.Onemoreisolated fragmentbelongstothisjaw.Theimportantmandiblemea surementsforsexidentificationarelistedinTable2forthe morecompleterightmandible.Theteethinbothupperand lower jaws are similarly unworn and unpolished. Vertebralcolumnandpelvis .…Manyofthe39vertebrae preserved(Fig.7;Table1)arealmostcomplete,butalso (#9) Thearticulatedandalmostcompleteskeleton3ofthesteppelion Pantheraleospelaea (Goldfuss,1810)fromtheSrbskoChlum-KomínCave (NMP No. R 4406). The pathologically damaged brain-case was in the process of healing (after Diedrich & ™ák 2006). !!"#$%&"'

PAGE 12

(#:) Craniumoftheyoungadultlioness Pantheraleospelaea (Goldfuss,1810)skeleton(skeleton3)fromtheSrbskoChlum-KomínCaveofthe BohemianKarst(CzechRepublic).€A…skullwithmandibledorsal ,B…skullwithmandiblelateralright ,C…skullwithmandibleventral ,D…skullven tral, E … skull frontal, F … mandible dorsal, G … mandible lateral right (NMP No. R 4406). (#;) Vertebralcolumnoftheyoungadultlioness Pantheraleospelaea (Goldfuss,1810)skeleton(skeleton3)fromtheSrbskoChlum-KomínCave oftheBohemianKarst(CzechRepublic). €A…firstcervicalvertebra,atlas(NMPNo.R4409) ,a…cranial,b…dorsal .€B… secondcervicalvertebra,axes (NMPNo.R4410),a…dorsal ,b…lateral. €C…thirdcervicalvertebra(NMPNo.R5032) ,a…cranial,b…dorsal .€D… fourthcervicalvertebra(NMP No.R4414) ,a…cranial ,b…dorsal .€E… fifthcervicalvertebra(NMPNo.R4411) ,a…cranial ,b…dorsal .€F… sixthcervicalvertebra(NMPNo.R4412), a…cranial ,b…dorsal .€G… seventhcervicalvertebra(NMPNo.R4413) ,a…cranial ,b…lateral .€H… firstthoracicvertebra(NMPNo .R4548), a…cranial, b…lateral .€I… secondthoracicvertebra(NMPNo.R4549) ,a…cranial ,b…lateral .€J… thirdthoracicvertebra(NMPNo.R4551) ,a…cranial ,b…lateral.+ + + + + +# +

PAGE 13

" €K…6…7thoracicvertebra(NMPNo.R4550),a…cranial,b…lateral .€L…7…8 thoracicvertebra(NMPNo.R4552),a…cranial,b…lateral. €M…ninth thoracicvertebra(NMPNo.R4553) ,a…cranial,b…lateral. €N…tenththoracicvertebra(NMPNo.R4555) ,a…cranial,b…lateral. €O…elevenththoracic vertebra(NMPNo.R4554),a…cranial,b…lateral.€P…twelfththoracicvertebra(NMPNo.R4556),a…cranial,b…lateral.€Q…thirteenththoracicver tebra(NMPNo.R4557) ,a…cranial ,b…lateral .€R… fourteenthandlastthoracicvertebra(NMPNo.R4558) ,a…cranial ,b…lateral .€S… firstlumbarver tebra(NMPNo.R4559) ,a…cranial ,b…lateral .€T… thirdlumbarvertebra(NMPNo.R4561) ,a…cranial ,b…lateral.€U…sixthlumbarvertebra(NMP No.R4563) ,a…cranial,b…lateral. €V…seventhandlastlumbarvertebra(NMPNo.R4564) ,a…cranial,b…lateral. €W…uppercaudalvertebra(NMP No.R4566),lateral .€X… uppercaudalvertebra(NMPNo.R4567),lateral .€Y… uppercaudalvertebra(NMPNo.R4568),lateral .€Z… uppercaudal vertebra(NMPNo.R4569),lateral.€AA…uppercaudalvertebra(NMPNo.R4570),lateral.€BB…middlecaudalvertebra(NMPNo.R4571),lateral. €CC…middlecaudalvertebra(NMPNo.R4597),lateral.€DD…middlecaudalvertebra(NMPNo.R4572),lateral.€EE…middlecaudalvertebra(NMP No.R4573),lateral.€FF…lowercaudalvertebra(NMPNo.R4574),lateral.€HH…lowercaudalvertebra(NMPNo.R5378),lateral.€II…lowercaudal vertebra (NMP No. R 5380), lateral. € JJ … last two fused lower caudal vertebrae (NMP No. Ra 4218), lateral.* 3!( + * 3 ! ( 55 44 66 2 7 78 8, , 2 << . . / == > ? @A 33!!(( B++** / 554466 + !!"#$%&"'

PAGE 14

includefragmentsoftheprocesses.Somevertebraestill havepartsmissingasaresultofdamagethatoccurreddu ringexcavation.Thecervicalcolumn(Fig.7A…G)iscom plete,withallsevenvertebraepresent,butmostofthese showrecentdamageontheirprocesses.Thethoracicver tebralcolumn(Fig.7H…R)isincomplete,asonly12ver tebraearepresent;thereshouldbe13intheFelidae.One vertebrafromthemiddlethoracicregionismissingand anotherisonlyrepresentedbythedorsalspine.Thelumbar vertebralcolumn(Fig.7S…V),whichinthefelidsconsists ofsevenvertebrae,ismissingthreelargevertebracentra,of whichmostoftheprocessesseemstobepresent.Eventhe halflumbarvertebra7ispresent.Fifteencaudalvertebrae (Fig.7W…JJ)arepresentfromthetail,includingthelasttwo fusedvertebraefromthetip.SinceacompletetailintheFe lidaehasabout22vertebrae,someareclearlymissing. Thethoraxincludesoneofthebestpreservedandmost completeribcagesknown(Fig.8A…Q).Someribsaremiss ingtheribheads,buttheirdistalendshaveatleastbeenpre served. P.l.spelaea has13ribsoneachsidecorrelatingto the13thoracicvertebrae.All26ribsarerepresentedinthis lionessskeleton,andaremoreorlesscomplete.Therightrib (#) Thoraciccostaeandsternalbonesoftheyoungadultlioness Pantheraleospelaea (Goldfuss,1810)skeleton(skeleton3)fromtheSrbsko Chlum-KomínCaveoftheBohemianKarst(CzechRepublic). €A…secondrightcosta(NMPNo.Ra4225) .€B… thirdrightcosta(NMPNo.Ra4226). €C…fourthrightcosta(NMPNo.Ra4227) .€D… fifthrightcosta(NMPNo.Ra4228) .€E… sixthrightcosta(NMPNo.Ra4248) .€F… ninthrightcosta (NMPNo.Ra4233) .€G… tenthrightcosta(NMPNo.Ra4234).€H…eleventhrightcosta(NMPNo.Ra4235) .€I… twelfthrightcosta(NMP No.Ra4236) .€J… thirteenthandlastrightcosta(NMPNo.Ra4237) .€K… firstleftcosta(NMPNo.R4238) .€L… secondleftcosta(NMPNo.Ra4239). €M…sixthleftcosta(NMPNo.Ra4243) .€N… seventhleftcosta(NMPNo.Ra4244) .€O… ninthleftcosta(NMPNo.Ra4246) .€P… tenthleftcosta (NMPNo.Ra4247).€Q…thirteenthandlastleftcosta(NMPNo.Ra4251).€R…firststernalbone(NMPNo.Ra4252).€S…middlesternalbone(NMP No. Ra 4253). All in inner lateral view. (#C) Appendicularskeletonoftheyoungadultlioness Pantheraleospelaea (Goldfuss,1810)skeletonfromtheSrbskoChlum-KomínCaveofthe BohemianKarst(CzechRepublic). €A…rightscapula(NMPNo.R4415),lateral. €B…leftscapula(NMPNo.Ra4233),lateral. €C…righthumerus (NMPNo.R4417),cranial .€D… lefthumerus(NMPNo.R4522),cranial .€E… rightulna(NMPNo.R4418),cranial .€F… leftulna(NMPNo.R4419), cranial. €G…rightradius(NMPNo.R4420),cranial. €H…leftradius(NMPNo.R4421),cranial. €I…rightpisiform(NMPNo.R4621),dorsal.€J…left pisiform(NMPNo.R4934),cranial. €K…leftscapholunatum(NMPNo.R4606),cranial. €L…pelvis ,a…dorsal ,b…lateralleft .€M… rightfemur* 3 ! 2 , 8 (5467< . +

PAGE 15

(NMPNo.R4525),cranial. €N…leftfemur(NMPNo.R4526),cranial. €O…incompleterightfibula(NMPNo.R4607),lateral. €P…incompleteleftfib ula(NMPNo.R4532),lateral. €Q…rightpatella(NMPNo.R4602),cranial. €R…leftpatella(NMPNo.R4533),cranial. €S…righttibia (NMPNo.R4528),cranial .€T… lefttibia(NMPNo.R4527),cranial .€U… leftcalcaneus(NMPNo.R4530),cranial .€V… rightastragal(NMP No. R 4914), dorsal. € W … left astragal (NMP No. R 4531), dorsal. € X … left tarsal II (NMP No. Ra 4232), cranial.* 3 ! 4 6 . 5 ( , 7 < > 2 @ = / ? 8 10 cm+ !!"#$%&"'

PAGE 16

cagehasthefollowingcompleteribs:Nos.2,5,6,and9…13. Oftheotherribs,Nos.1,3,and4havesmallpartsmissing, No.7hasnodistalpart,andNo.8hasnoproximalribhead. Intheleftribcagetheribs1…2,6,9and13arecomplete;all othershavesomedistalparts(Nos.3…5)ortheproximalpart (No.12)missingor,inafewcases,minordamageinthe centralpartoftheribshaft(Nos.8,10).Thefirststernal boneispresentaswellasoneotherwhichcouldbeeitherthe secondorthirdsternalbone(Fig.8R,S). Thepelvisisalmostcomplete(Fig.9L)butisstillmissingalargeportionoftheposteriorpart.Allbonesutures, excepttheonesbetweenthepelvicbonesandsacrum,are fused, indicating a young adult age. Appendicularskeleton .…Theforelimbs(Fig.9A…K)arealmostfullypreserved.Onlythedistalelementsareproblematic, i.e .thelastphalanges(II…III).Theleftscapulaisalmostcompletewhiletherightscapulaislesscomplete (Fig.9A,B).Therighthumerushasasmallfragmentmis sing,whiletheleftoneiscomplete(Fig.9C,D).Theright ulna(Fig.9E)iscomplete,asisalsotheleftone(Fig.9F). Bothradiihavesimilarproportions:theleftone(Fig.9H) hasalargefragmentmissingbuttherightoneiscomplete (Fig.9G).Bothcompletepisiformbonesarepresent (Fig.9I,J).Therightscapholunatumispresent(Fig.8K). Halfofthemetacarpalsaremissing.Allpedalboneshave beensortedfromthenearlycompletemanusandpesmeta podialsetsofthejuvenileanimal,whichareagainsmaller andcontainallMcII…Voftherightandleftmanus.Only therightMcIII…VandtheleftMcI,IV…Vseemtobepre sentfromtheadultfemaleskeleton.AtleastonephalanxI wasseparatedfromthejuvenilematerial.Onlythreepha langesfromtherightmanusandthreefromtheleftmanus canprobablybeattributedtothelionessskeleton.Theori ginofphalanxIIwasmuchmoredifficulttodistinguish anditisevendifficulttodeterminearightorleftposition. TheexactpositionsofthephalanxII,andalsothephalanx III,havethereforenotyetbeenidentified.Theymayeven belongtothepedalskeletons,orhavederivedfromthecub skeleton. Bothhindlimbbones(Fig.9M…X)arealmostcom plete.Bothfemora,thepatellaeandthetibiaeareonly partlycomplete(Fig.9S,T).Onlyhalfofthefibulaeare present.Theleftcalcaneus(Fig.9U)ispresentbutthe rightoneisabsent.Bothastragals(Fig.9V,W)arepre servedandarelargerthenthosefromthejuvenileanimal. Twolefttarsals(cuboid,navicular)arecomplete,whereas onlyhalfofthetarsalIIispresent(Fig.9X).Theleftpes includesalllargemetatarsiII…V.Fromthisleftpestwo morefirstphalanges(digitIV,anddigit?),onesecond phalanx(digitII),andpossiblyallthirdphalangesare present.Thereisagainaproblemintheattributionofan exactpositiontothispartoftheskeleton.Thephalanges IIIcanbedistinguishedbetweentherightandtheleftside asaresultoftheirdifferentanglestothephalanxbases. Theexactpositionofthedigitsisalsouncertain,dueat leastinparttotheirbeingincomplete.SomesmallphalanxIII,whichhavebeenattributedtotheouterdigits, mayactuallycamefromtothemiddledigitsofthesmaller juvenileanimal.Therightpedalskeletonhasonlythree completelargemetatarsi(V,IV,andII)andhalfofthe proximalmetatarsusII.Thefirstphalanxispresentfrom thedigitsII,IIIandV.PhalanxIImayberepresentedon digitsII…IV,butthisisuncertain.Finally,thepositionof thephalangesIIIinrelationtodigitsIIandVremainsun clear.3 "####!ThesexofthelionsfromtheCzechRepubliccanbeinter pretedthroughcomparisonswithotherfossilsoffemale andmalelions( cf .Turner1984,Gross1992,Diedrich 2009b).Femalesaremuchsmallerthenmales,asdefined differentlyforcoldandwarmperiods,usingosteometrics forthecraniumandpostcranialbones(Fig.11)withincold andwarmperiods,butnotbetweenthosetimes.Thelioness skeletonfromtheEemiansiteatNeumark-NordLake1 / -) Importantmeasurementsofthe Pantheraleospelaea skeletonfromSrbskoChlum-KomínoftheBohemianKarst(CzechRepublic)forthesex identification (in cm). The measurements show low proportions and indicate well a female individual (all in mm). 1R 4406CraniumTotal length: 305165Width P4: 34Width C: 21 2R 4407Lower jawTotal length: 214Height behind M1: 48Width M1: 28Width C: 21 3R 4522Humerus (left)Total length: 319Smallest width middle of shaft: 29Proximal width: 75Distal width: 84 4R 4419Ulna (left)Total length: 342Width under proximal joint: 48Proximal width: 60Distal width: 34 5R 4421Radius (left)Total length: 288Smallest width middle of the shaft: 29Proximal width: 39Distal width: 62 6R 4523PelvicTotal length: 312Width “canal”: 92Acetabular width: 132Sacrum length: 102 Acetabulum length: 42 Length foramen obturatum: 77 7R 4525FemurTotal length: 359Smallest width middle of the shaft: 34Proximal width: 88Distal width: 81 8R 4528TibiaTotal length: 312Smallest width middle of the shaft: 29Proximal width: 84Distal width: 59

PAGE 17

( cf .Diedrich2010b,c)andasolitaryskullfromanItalian cave(Bona2006)werenotincludedinthestatisticsbe causesteppelionsseemtohavebeensmallerduringwarm periods(includingthepresent)thanincoldperiods( cf . Gross1992),althoughthisisnotyetwelldocumentedin thefossilrecordbecauseofthescarcityofEemianlionma terial in Europe. Skeletons1and2andskullremainsfromtheSloupCave.– Bothskullsofskeletons1and2fromtheSloupCavefall (#) AllrecentlydiscoveredLatePleistocenesteppelion Pantheraleospelaea (Goldfuss,1810)craniafromtheCzechRepublic .€A… skullfrom theopenairloessBerounkaRiverterraceHýskovsitenearBeroun(MBKBNo.363a) .€B… skullwithlowerjawfromskeleton3,theyoungadultlioness skeletonfromtheSrbskoChlum-KomínCave(NHMPNo.R4406).€C…skullwithlowerjawfromtheCut-StoneGalleryoftheSloupCavecomposite skeleton2(AMBwithoutNo.).€D…skullfromtheCut-StoneGalleryoftheSloupCave(AMBNo.OK130570). €E…skullwithlowerjawfromthe Cut-Stone Gallery of the Sloup Cave composite skeleton 1 (NHMV No. 1885/0014/4302). All in dorsal view.* 3! + !!"#$%&"'

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withintherangeofthosefromsmalltomedium-sizedma les(Fig.11A).ThethirdLatePleistoceneskullfromthe SloupCave,whichisthelargestoftheisolatedCzechskulls (Fig.10D,totallength378mm,condyluswidth72mm),is intermediateinsizebetweenskulls1and2.Maleskulls fromSiegsdorfandAzéareevenlarger(withlengthsbe tween380and420mm, cf .Gross1992;Fig.11),asisthe skullofthelargemaleskeletonfromArrikrutz(407mm, cf . Altuna1981).ThelongbonesofboththeSloupCaveskele tonsaredissimilarinlength(exceptthehumeriinskeleton 1)butarelessmixedinskeleton1(Vienna),whichconsists mainlyofmalebones( cf .Fig.11B…F).Skeleton2(Brno)is amixtureofmaleandfemaleremains(Fig.3B)withsignifi cantvariationsinlongbonedimensions( cf .Fig.11B…F). Skeleton3(SrbskoChlum-KomínCave).– Thetotalskull lengthofthisspecimenisabout302mmwhichisthesame asthatoftheadultfemaleskullfromthePerickCavesin Germany(Fig.12G;Diedrich2007b).However,skeleton3 appearstobefromayounganimalwhoseskullproportions aresimilartotheimmaturelionessskullfromtheUrilor Cave( cf .Fig.12A,D).Amoreusefulindicatorforsex identificationisthesmallupperjawP4(PerickCaves: 39mm;SrbskoChlum-Komín:34mm;Siegsdorf:40mm; Arrikrutz:43mm).Finally,thelowerjawsaresmallerthan thosefromSiegsdorfandArrikrutz.Themandibleheight behindtheM1fortheSrbskoChlum-Komínspecimenis only48mm,whereasforSiegsdorfitis55mmandforArrikrutz60mm,whichwhichsuggeststhatitisinthefemale sizerange.Measurementsfromskull3areclearlyinconclusive with regard to the age or sex of this specimen. Similarvariationsindimensionsareapparentbetween thelongbonesfromtheSrbskoChlum-Komínlionessand thosefromthemalelionskeletonsofSiegsdorfand Arrikrutz(seeFig.11).TheSrbskoChlum-Komínspeci menisalargefemale(oryoungmale),whilethelionfrom Siegsdorfisamediumsizedmaleandthelionfrom Arrikrutz is a very large male. ThelionessfromtheSrbskoChlum-KomínCaveisthe onlyimmaturelionmaterialfromthatcaveand99%ofthe steppelionmaterialfromtheSloupCavecanbeattributed toadultanimals.Asimilarsituationoccursinalmostallof theEuropeancavebeardensinwhichlionremainshave beenfound,supportingtheclaimthatthesecaveswerenot usedas“caveliondens”( cf .PerickCaves,BilsteinCaves: Diedrich 2009a, b, Urilor Cave: Diedrich et al . 2009).$%#!% &#' #'Threeskulls,fromtheSrbskoChlum-Komín,Sloupand Zoolithencaves,exhibitdamagethatoccurredwhilethe animalwasstillalive(Fig.12A…C).Allshowdamageto thebrain-caseandsaggitalcrestthatwasintheprocessof healing.Theobservedcallusformationanddeformation structureswereprobablyareactiontobitedamage.One longtoothscratchmarkisclearlyvisibleontheZoolithen Caveholotypeskull(Fig.12C).Thebrain-caseoftheSrb skoChlum-Komínlionexhibitsatraumaontheleftside (Fig.12B)affectingtheparietalboneandleavinganelon gateddepressionwithaparallelcrest.Thisfeaturerunspa ralleltothecristasaggitalisoverthecentreoftheparietal bone.Thisdamagealsocausedamarkeddeformationof theleftsideofthebrain-caseandresultedincompletefu sionofthesuturebetweentheleftparietalandthefrontal bones.Theparietalfractureorscratchmayhaveaffected theentireleftsideofthebrainincludingmotorfunction, auditory,andsensoryareas.Theskullfromskeleton1 (Vienna)alsoseemstohavebitedamagethatcausedminor deformationontheparietal.Thiswasalmostcompletely healedbythetimeofdeath.Frontaldamagetoacavebear skullfromtheZoolithenCaveisalsopartlyhealed (Fig.13C).Otherbrain-casepathologiesona P.l.spelaea craniumfromtheopenairsiteatHaltern,north-western Germany,thatoccurredwhiletheanimalwasstillalive, havebeendescribedandillustratedpreviously(Diedrich 2004).Thisskullshowsalargebonegrowth1cmindiameterontheparietal,closetothesaggitalcrest,whichhasnot yetbeenexplained.Damageandpathologiesreportedin modernAfrican P.leo (suchaspartialcerebellahernia… Tuch&Pohlenz1973),havenotbeenobservedinLate Pleistocenesteppelionskulls,andappeardifferenttothe bitedamageandpathologiesonPleistocenecranialmaterialdiscussedherein.Othertypesofdepressionsinparietals,occurringintheLatePleistocenesabretoothcat Smi lodonfatalis ( e.g .,deeppitsbelievedtobeneoplasms), havebeeninterpretedtobetheresultofmechanicalstrain (Duckler1997).Theyarenotcomparabletotheparietal damagepresentedonthethreesteppelionskullsof Fig.12A…C,whicharebelievedtohavehadtraumaticcau sesandtomostprobablyresultfrombitedamagecausedby canine teeth during fights. Intraspecificfightsbetweenpresent-dayfemalelionsor betweenmaleandfemalelionsarequitecommonandocca sionallyresultinthedeathofoneofthecombatants ( cf .Palomares&Caro1999,Packer&Pusey2001).Fights inwhichlionsarecriticallyinjuredorkilledmayoccurdur inganattemptedtakeoveroftheclanbyamalelionorwhen femalesvigorouslydefendtheircubs(Estes1999,Packer& Pusey2001).Fightinglionsoftenbiteeachotherintheface andhead(Schaller1972,Packer&Pusey2001).Suchsce narioscouldexplaintheparietalinjuryandbraindamagein thePleistoceneearlyadultfemalelionskeleton(Fig.5)from SrbskoChlum-KomínandtheskullsfromtheSloupand Zoolithencaves(Fig.12A…C),andwouldalsoexplainthe brain-casedamageofotherLatePleistocenesteppelions.

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Lionsarealsoknowntofightforotherreasons,includ ingdisputeswhilefeedingonacarcass,territorialdisputes betweenprides,andfightsbetweengroupsofmales,which canallresultinsimilarbitewounds(Schaller1972,Estes 1999, Packer & Pusey 2001). Interspecificfightsalsooccurbetweenpresent-day# (#) LatePleistoceneWeichselian/Wuermiansteppelion Pantheraleospelaea (Goldfuss,1810)sexualdimorphism:comparisonsofskulland longbonedimensionsforCzechRepublicmaterialwiththosefromotherEuropeansites(composedafterAltuna1981;Argant1988;Diedrich2009b, 2011a; Diedrich & Rathgeber 2011). !!"#$%&"'

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lionsandhyenas,with C.c.crocuta beingoneofthemain predatorsinAfrica.Thereareseveralanimalspeciesthat cankillhyenas,butlionsaretheirmainkillers( cf .Joubert &Joubert2003).Inonestudy13outof24hyenacarcasses foundhadbeenkilledbylions(Kruuk1972,Joubert& Joubert2003)andasimilarsituationislikelytohaveex istedwiththeLatePleistocenelionsandIceAgespotted hyenas.Possiblesupportingevidencecomesfromhyena skullfromtheZoolithenCave(Fig.13A)whichexhibits considerablebitedamagethathadpartlyhealed,although itcannotyetbeaccuratelydeterminedfromthefossilre cordwhetherspecificinjurieswereconspecificorinflicted byanotherspecies.Similarlyforattacksonthesteppelion skullfromthesamecave(Fig.13B).Actualisticcompari sonswillneedtobemadeinfutureonpresentdayhyenas andlionstocomparethedifferenttypesofdamageinflicted bytheirbites.Wherelionsareseparatedfromtheirprideor arenotintegratedintoapride,individualAfricanlionsdo feedoncarcassesinordertosurviveinemergencies(Estes 1999).IftheinjuredyounglionessfromtheSrbsko Chlum-KomínCavetriedtointrudeintoahyenaden,the chancesofitwinningabattlewithhyenasdefendingtheir prey-storageandcub-raisingdensitewouldbeextremely low.Suchadefenceofdensandjuvenilesprovidestherea sonformostofthelionkillsbypresent-dayhyenasinAf rica(Estes1999,Packer&Pusey2001).Incontrast,Africanspottedhyenasarewellknowntokilllioncubs, juveniles,andweakorsickadultlions(Schaller1972, Estes1999).Consumptionofthevictimappearstobemore commonwhenfoodisscarceoramatterofdispute (Palomares&Caro1999),whichexplainsthepresenceof completePleistocenelioncarcassesinhyenaorcavebear den sites. Interspecificfightsbetweenlionsandcavebearshave nomodernequivalentand,possiblyasaresult,the taphonomyofcavebeardencavesinEuropehasnotbeen wellstudied.Thetheorythatsteppelionsmayhavehunted cavebearsduringtheIceAgewasestablishedonthebasis ofnewdiscoveriesattheUrilorCaveandothersitesin GermanyandtheCzechRepublic( cf .Diedrich2009b, 2010e,2011f).Afirststudyofthecavebearbone taphonomyintheZoolithenCave(Diedrich2011f)has suggestednewexplanationsforpartlyhealedbitemarkson thebrain-cases(frontals)ofcavebears( cf .Fig.13C).Only intheZoolithenCavehavesuchimpressivelydamaged skullsofthetwolargecaveinhabitants(hyenasandcave bears)andothercavedwellers(lions)beenfound,prompt ingthetheorythathyenasscavengedoncavebearsandthat thecavebearswereactuallyhuntedbylions(Diedrich 2010e,2011f).Thecranialdamageinflictedonlivingcave! (#-) Premortempathologiesandpostmortemskulldamageofthesteppelion Pantheraleospelaea (Goldfuss,1810) .€A… skeleton3skullfrom theSrbskoChlum-KomínCavewithbitedamageonthebrain-case,intheprocessofhealing .€B… skeleton1skullfromtheSloupCavewithbitedamage intheprocessofhealingandskulldeformation. €C…holotypeskullfromtheZoolithenCavewithbiteandscratchdamageonthesaggitalcrestofthe brain-case. €D…scavengedskullfromtheUrilorCaveskeletonwithbitedamageontheoccipital.€E…skullfromtheZoolithenCavewithbrain-case openingandfracturedjugal .€F… lowerjawfromthePerickCaves,fracturedtoremoveitfromtheskull .€G… skullfromthePerickCaveswithjugal arches fractured for lower jaw removal.* 3 + ( !

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bearscouldalsobepossiblyattributabletointraspecificaggression.Itisfurtherpostulatedthatbattlesbetweencave bearsandlionsmayhaveoccurreddeepwithinthecave,in areasthatwouldhavebeeninaccessibletohyenasdueto theirrelativelypoorclimbingskills( e.g .UrilorCave, Diedrich2011f).Cavebearsmaythushavedeliberately soughtdeepcavesinwhichtohibernate,asprotection against lion predation (Diedrich 2009b, 2011f).$%#! #&'!!Someofthebitedamageonlionskullsisobviouslypost mortemdamage,includingthatseenonthethreeskulls fromtheUrilor,ZoolithenandPerickcaves(Fig.12D…G). Thebitemarksshownosignofhealingandaretypically roundorovaltotriangular,manywithassociatedscratch marks.Insomeinstancesportionsoftheboneshavebeen chewedoff.IntheZoolithenCaveskull,thebrain-case musthavebeenopenedfromtheoccipitalsidebyalarge carnivoreasamissingjugalarchhasalsobeenchewed. Jugalarchdamageisalsodemonstratedonbothsidesofthe skullfromthePerickCaves(Fig.12G).Thelowerjaw fromthisskullhasalsobeenfractureddiagonallybehind thelastmolartoothinordertofacilitateitsremovalfrom theskull(Fig.12F).ThepredationofIceAgespotted hyenaonlionsandimportationoftheircarcasesintohyena cavedensmayprovideanexplanationforthegeneral male/femaleratiosinlatePleistocenebonesites( cf .Died rich2009a).Modernspottedhyenasdonotusuallyhunt adultmalelions,althoughexceptionstothisgenerality havebeenreported(Kruuk1972,Estes1999).Somecar cassesofIceAgesteppelionsmayhavebeenleftoutside thecaves(orpartlyimportedtohyenadens),asinPrahaPodbabariverterraceopenairsiteandattheSiegsdorf openairriverterraceandlakesiteinGermany,although theimportationofmalelioncarcasseshasnowbeende monstratedinsomeEuropeanhyenadens( cf .Diedrich 2007c,2009a).Thereisacorrelationbetweenthemoreda magedlionbonesandtheiroccurrenceathyenadensites andprey-boneaccumulations.Wheretherewerenohyenas occupyingtheentrancesorside-branchesofcavebearden cavesystems( cf .Fig.13)thedamagetoskeletonsisredu ced,ashasrecentlybeenshownfortheUrilorCave(Died rich2011f).IntheZoolithenCave,thelargehyenapopula tionwasalsothoughttohavebeenresponsibleforthehigh proportionanddegreeofdamagedcavebearbones,ashas alsobeendescribedforthePerickCavesandothercave beardensitesinEurope( cf .Diedrich2011f).Thebite (#0) CavemodelfortheSloupCave,ZoolithenCave,andotherlargeEuropeancavesthatwereusedashyenaandcavebeardens.€A…C…three skullsfromtheZoolithenCaveallhavepartlyhealedbitedamageonthebrain-case(materialintheMBandUE).Suchdamagecouldhaveresultedfrom differentscenariossuchasconflictsbetweenhyenasorlionsandtheircavebearprey,orfrominterspeciesfights,eitherinsideoroutsidethecave .Lion skeletonsfromcavebeardens,suchasthosefromtheSloupCave,areallfounddeepwithinthecaveswhereasincompleteandgnawedlionbonesinthe hyena den area appear in several cases to have been imported lion remains.* + !!"#$%&"'

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damageleftbyhyenasoncavebearbonesissimilartothat illustratedhereinonlionremains.Thelionskullsfromthe PerickCaves(Fig.12F,G)…aswellasremainsfromthe SloupCave( cf .Diedrich2011b)…thathavepossiblybeen chewedandcrackedbyhyenasexhibittypicaldamageto thejugalarchescausedduringremovalofthelowerjaw, whichalsoresultedincrackingofthemandibles.While hyenasseemtohavebeenthemainscavengersonlioncar cassesit remainsapossibilitythat,instressfulsituations, lionsthatperhapsbecametrappedinacomplexcavesys temwithseveraldifferentlevelsmighthaveengagedin cannibalism ( e.g . Urilor Cave: Diedrich 2011f).ThehistoricalskeletonfindsfromtheCut-StoneGalleryin theSloupCaveoftheMoravianKarst(CzechRepublic) arebothcompositeskeletons.Skeleton1(Vienna)includes ahigherproportionoforiginalbones,whichhavecome fromasinglemaleskeleton.Bothofthesecompositeskele tonsincludeabout30%to50%bonecastsandalsopedal bonesfromtheVýpustekCaveintheCzechRepublic.The lionbonesandskeletalremainswerefoundeitherwithin hyenadensandprey-boneaccumulations,ordeeperwithin cavebeardens.ThematerialfromtheSloupCaveindicates derivationfromseveraldifferentsteppelion Pantheraleo spelaea (Goldfuss,1810)skeletons,foundwithinthecave bearhibernationareasdeepinsidethecave,withallofthe materialbeingfromadultlionsorlionesses.Thissituation isdifferentfromtherelativelycompleteoriginalskeleton ofadiseased,youngadultlionessfromtheSrbsko Chlum-KomínCavehyenadensiteintheBohemianKarst (CzechRepublic),whichmayhavebeenkilledbythe hyenaclanthatoccupiedthesamecave,whenitattempted tostealtheirprey.Inoverlappinghyenaandcavebeardens suchastheSloupCaveandtheGermanZoolithenCave, thelionbonetaphonomyismorecomplex.Hyenasappear tohavedispersedthelionskeletonsandmayalsohaveim portedpartsofsomelioncarcassesforconsumptioninside thecave.Insodoing,theydamagedbones,opened brain-casesinordertofeedonthebrain,andfractured thelowerjaws.Injuriesonsteppelionskullsfromdifferent caves,thatwereintheprocessofhealing,provideevidence ofeitherintraspecificconflictsbetweenlionsorinterspeci ficfightsbetweenlionsandeitherhyenasorcavebears ( e.g .,partlyhealedcranialbitedamage,ZoolithenCave (Fig.12).Thepresenceofarticulatedlionskeletonsdeep insidecavebeardenscanbeexplainedbyinterspecific fightsduringpredationbylionsoncavebears,inwhichli onswerekilledbybears.Lessthen1%ofthebonesincave beardensarefromsteppelionsbutitappearsthat,from timetotime,alionintentonpredationmayhavebeenkil ledbythecavebearsprotectingtheircubs.Cavebearswere notcarnivorousandhencewouldnothavefedonanylions thattheykilled,andhyenasoftenhadnoaccesstothedee perpassagesinthecaves.Thepresenceofsteppelionske letonssuchasthethreerecentlydiscoveredbetweenarticu latedcavebearskeletonsandbonebedsonhibernation plateausdeepwithintheRomanianUrilorCave,appearto offerataphonomicmodelforthepresenceofsteppelion carcassesandbonesinotherlargecavebeardenssuchas theSloupCave,theVýpustekCave,theZoolithenCave, the Perick Caves, and many other European sites.+%#ThecollectionintheNaturalHistoryMuseum,Vienna,wasstud iedwiththesupportofU.Göhlich,andthecollectionintheNa tionalMuseum,PraguewiththesupportofK.Zágorek.Avisitto inspectthecollectionsintheAnthroposMuseum,Brno,waspos siblywithpermitfromM.Oliva.R.Musil(UniversityofBrno) actedasguideintheSloupandVýpustekcavesandalsoprovided accesstorelevantliterature.O.Hampe(MuseumofNaturalHis toryoftheHumboldtUniversityBerlin)andB.Hilpert(Univer sityofErlangen)allowedthestudyofZoolithenCavematerial. TheFHKFspeleologicalclubassistedwiththecavefieldwork andwithnewexplorationandresearchinthespringandsummer of2010.ThefirstdraftwaskindlyreviewedandcorrectedbyR. MusilandP.A.White.Thefinalspell-checkandcommentswere thankfully made by E. Manning.ALTUNA,J.1981.FundeinesSkelettesdesHöhlenlöwen ( Pantheraleospelaea Goldfuss)inArrikrutz,Baskenland. Bonner zoologische Beiträge 32(1–2) , 31…46. ARGANT,A.1988.Étudedel’exemplairede Pantheraspelaea (Goldfuss,1810)(Mammalia,Carnivora,Felidae)dugise mentPleistoc è nemoyenrecentdelagrotted’Aze(Saoneet Loire). Revue de Paléobiologie 7(2) , 449…466. BONA,F.2006.Systematicpositionofacompletelion-likecat skullfromtheEemianossiferousrubblenearZandobbio(Ber gamo,NorthItaly). RivistaItalianadiPaleontologiaeStra tigrafia 112(1) , 157…166. BURGER,J.,ROSENDAHL,W.,LOREILLE,O.,HEMMER,H.,ERIKSSON, T.,GÖTHERSTRÖM,A.,HILLER,J.,COLLINS,M.J.,WESS,M.J.T. &ALT,K.W.2004.Molecularphylogenyoftheextinctcave lion Pantheraleospelaea . MolecularPhylogenyandEvolu tion 30 , 841…849. DOI 10.1016/j.ympev.2003.07.020 DIEDRICH,C.2007a.UpperPleistocene Pantheraleospelaea (Goldfuss,1810)skeletonremainsfromPraha-Podbabaand otherlionfindsfromloessandriverterracesitesinCentralBo hemia(CzechRepublic). BulletinofGeosciences82(2) , 99…117. DOI 10.3140/bull.geosci.2007.02.99 DIEDRICH,C.2007b.Thefairytaleaboutthe“cavelions” Pantheraleospelaea (Goldfuss1810)ofEurope…LateIce AgespottedhyenasandIceAgesteppelionsinconflicts…lion killersandscavengersaroundPrague(CentralBohemia).

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ScriptaFacultatisScientiarumUniversitatisMasarykianae Geology35(2005) , 107…112. DIEDRICH,C.2008.TheholotypesoftheUpperPleistocene Crocutacrocutaspelaea (Goldfuss,1823:Hyaenidae)and Pantheraleospelaea (Goldfuss,1810:Felidae)ofthe ZoolithenCavehyenaden(SouthGermany)andtheir palaeo-ecologicalinterpretations. ZoologicalJournalofthe Linnaean Society London 154 , 822…831. DIEDRICH,C.2009a.SteppelionremainsimportedbyIceAge spottedhyenasintotheLatePleistocenePerickCaveshyena deninNorthernGermany. QuaternaryResearch71(3) , 361…374. DOI10.1016/j.yqres.2008.12.006 DIEDRICH,C.2009b.UpperPleistocene Pantheraleospelaea (Goldfuss,1810)remainsfromtheBilsteinCaves(Sauerland Karst)andcontributiontothesteppeliontaphonomy,palaeo biologyandsexualdimorphism. AnnalesdePaléontologie 95(2009) , 117…138. DOI 10.1016/j.annpal.2009.07.001 DIEDRICH,C.2009c.ALatePleistocenewolverine Gulogulo (Linné,1758)skeletonremainfromtheSloupCaveinthe MoravianKarst,CzechRepublic. AnnalendesNaturhistori schen Museums Wien 110A , 123…132. DIEDRICH,C.2010a.SpecializedhorsekillersinEurope…foetal horseremainsintheLatePleistoceneSrbskoChlum-Komín CavehyenadenintheBohemianKarst(CzechRepublic)and actualisticcomparisonstomodernAfricanspottedhyenasas zebrahunters. QuaternaryInternational220(1–2) ,174…187. DOI 10.1016/j.quaint.2010.01.023 DIEDRICH,C.2010b.EinSkeletteinerkrankenLöwin Panthera leospelaea (Goldfuss1810)undanderemännlicheLöwenresteausNeumark-Nord. ArchäologieinSachsen-Anhalt Sonderband , 432…441. DIEDRICH,C.2010c.Adiseased Pantheraleospelaea (Goldfuss 1810)lionessfromaforestelephantgraveyardintheLate Pleistocene(Eemian)interglaciallakeatNeumark-Nord,central Germany. Historical Biology 2010(4) , 1…23. DOI 10.1080/08912963.2010.507814 DIEDRICH,C.2010d. TheLatePleistocenesteppelionPanthera leospelaea(Goldfuss1810)populationfromtheZoolithen Cave(Bavaria)Germany–cavebearpredationadaptationin middlemountainoustaigaforestenvironmentsofCentralEu rope .Abstract,16thinternationalcavebearsymposiumAzé, France. DIEDRICH,C.2010e. Theholotypeskullof“Ursusspelaeus Rosenmüller1794”–insighttothecavebearbonetaphonomy intheZoolithenCave(Bavaria)Germany–andcontribution totheexplorationhistory,cavegenesis,andsedimentology . Abstract,16thinternationalcavebearsymposiumAzé,France. DIEDRICH,C.2011a.Thelargeststeppelion Pantheraleospelaea (Goldfuss1810)populationfromtheZoolithenCave(Ba varia)Germany:specialisedcavebearpredatorsofEurope. Historical Biology 23(2–3) , 271…311. DOI 10.1080/08912963.2010.546529 DIEDRICH,C.2011b.TheIceAgespotted Crocutacrocuta spelaea (Goldfuss1823)population,theirexcrementsand preyfromtheLatePleistocenehyenadenSloupCaveinthe MoravianKarst,CzechRepublic. HistoricalBiology (in press). DOI 10.1080/08912963.2011.591491 DIEDRICH,C.2011c.ALatePleistocenelion Pantheraleospelaea (Goldfuss1810)skullandotherpostcranialremainsfromthe SloupCaveintheMoravianKarst,CzechRepublic. Acta Archaeologica2011 , 19…28. DIEDRICH,C.2011d.LatePleistocenesteppelion Pantheraleo spelaea (Goldfuss,1810)footprintsandboneremainsfrom openairsitesinnorthernGermany…Evidenceofhyena-lion antagonismandscavenginginEurope. QuaternaryScience Reviews 30(15–16), 1883…1906. DOI 10.1016/j.quascirev.2011.03.006 DIEDRICH,C.2011e.Pleistocene Pantheraleospelaea (Goldfuss 1810)remainsfromtheBalveCave(NWGermany)…acave bear,hyenadenandMiddlePalaeolithichumancave,andre viewoftheSauerlandKarstlionsites. Quaternaire22(2) , 105…127. DIEDRICH,C.2011f.CavebearkillersandscavengersintheFinal IceAgeofEurope…feedingspecializationasreactiononthe mammothsteppefaunaabsenceinmountainousregions. Qua ternary International (in press). DIEDRICH,C.&RATHGEBER,T.2011.LatePleistocenesteppelion Pantheraleospelaea (Goldfuss1810)skeletonremainsofthe UpperRhinevalley(SWGermany)andcontributiontotheir sexualdimorphism,taphonomyandhabitus. HistoricalBiol ogy (in press). DOI 10.1080/08912963.2010.549943 DIEDRICH,C.,ROBU,M.,DRAGUSIN,V.,CONSTANTIN,S.&MOL DOVAN,O.2009. NewUpperPleistocenesteppelionskeleton findsbetweenthecavebearhibernationplateausofthe UrsilorCavebearden,Romania .Abstract,15thInternational Cave Bear Symposium, Spiská Nová Ves, Slovakia. DIEDRICH,C.&™ÁK,K.2006.UpperPleistocenehyena Crocuta crocutaspelaea (Goldfuss,1823)preydepositanddensitesin horizontalandverticalcavesoftheBohemianKarst(Czech Republic). Bulletin of Geosciences 81(4) , 237…276. DUCKLER,G.1997.Parietaldepressionsinskullsoftheextinctsaber-toothedfelid Smilodonfatalis :evidenceofmechanical strain. Journal of Vertebrate Paleontology 17(3) , 600…609. ESTES,R.1999. TheSafariCompanion:AGuidetoWatchingAf ricanMammals .ChelseaGreenPublishingCompany,Ver mont. GROSS,C.1992. DasSkelettdesHöhlenlöwen(Pantheraleo spelaeaGoldfuss1810)ausSiegsdorf/Ldkr.Traunsteinim VergleichmitanderenFundenausDeutschlandundden Niederlanden .UnpublishedDissertation,Tierärztliche Fakultät der Maximilians-Universität, München. GOLDFUSS,G.A.1810. DieUmgebungenvonMuggendorf.Ein TaschenbuchfürFreundederNaturundAltertumskunde . Erlangen. JOUBERT,D.&JOUBERT,B.2003. EternalEnemies:LionsandHy enas .WildlifeFilmsBotswanaforNationalGeographic,DVD. KAFKA,J.1903.FossileundRecenteRaubthiereBöhmens(Car nivora). ArchivderNaturwissenschaftlichenLandesdurch forschung von Böhmem 10(6) , 1…120. KRUUK,H.1972. Thespottedhyena.Astoryofpredationandso cial behavior . University of Chicago Press, Chicago. MUSIL,R.1956.MährischeFundstellenPleistozänerWirbeltiere. Geologie 5 , 159…179. MUSIL,R.2002. Sloupsko-šošvskéjeskyn .178pp.Publishing house Gloria, Brno. 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