Invertebrate fossils from cave sediments: a new proxy for pre-Quaternary paleoenvironments


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Invertebrate fossils from cave sediments: a new proxy for pre-Quaternary paleoenvironments

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Invertebrate fossils from cave sediments: a new proxy for pre-Quaternary paleoenvironments
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Biogeosciences
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Moldovan, O. T.
Mihevc, A.
Miko, L.
Constantin, S.
Meleg, I. N.
Petculescu, A.
Bosak, P.
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Sediments ( local )
Interior Caves ( local )
Interior Cave Facies ( local )
Fossils ( local )
Paleoenvironments ( local )
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serial ( sobekcm )

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Five samples of clastic sediments from interior cave facies taken in three Slovenian relic caves (Trhlovca, Račiška pečina, and a cave in Črnotiče Quarry, Classical Karst, SW Slovenia) provided invertebrate fossil remains. Most of them belong to Oribatida but sparse individuals of Cladocera and insects were also identified. They represent the first pre-Quaternary invertebrate fossils found in sediments of continental temperate climate. The Pliocene/Pleistocene age of the sediments was determined by paleomagnetic dating chronologically calibrated by micromammal biostratigraphy. Invertebrate fossils could be validated as new proxy for the study of cave sediments due to their suitability for ecological and paleogeographic correlations in caves and outside the caves. They also bring additional information about cave formation and karst hydraulic regime in the area. Although the number of remains was very low, it is evidence that climatic conditions in caves allow a better preservation of fossil remains of some groups as compared to most of the surface habitats. This may open a new direction in the study of cave sediments.
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Biogeosciences, Vol. 8 (2011-07-11).

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Biogeosciences,8,1825– 1837 ,2011www.biogeosciences.net/8/1825/2011/doi:10.5194/bg-8-1825-2011Authors2011.CCAttribution3.0License. Biogeosciences Invertebratefossilsfromcavesediments:anewproxyforpre-Quaternarypaleoenvironments O.T.Moldovan1,A.Mihevc2,L.Miko3,S.Constantin4,I.N.Meleg1,A.Petculescu4,andP.Bosak2,51DepartmentofCluj,“EmilRacovita”InstituteofSpeleology,Clinicilor5,400006Cluj-Napoca,Romania2KarstResearchInstitute,SRCSASA,Titovtrg.2,Postojna,Slovenia3EuropeanCommission,DGEnvironment,Av.deBeaulieu5,1160Auderghem,Brussels,Belgium4“EmilRacovita”InstituteofSpeleology,Frumoasa31,010986Bucuresti,Romania5InstituteofGeologyASCR,v.v.i.,Rozvojova269,16500Praha6,CzechRepublicReceived:18March2011–PublishedinBiogeosciencesDiscuss.:30March2011Revised:14June2011–Accepted:24June2011–Published:11July2011 Abstract.FivesamplesofclasticsedimentsfrominteriorcavefaciestakeninthreeSlovenianreliccavesTrhlovca,Raciskapecina,andacaveinCrnoticeQuarry,Classi-calKarst,SWSloveniaprovidedinvertebratefossilre-mains.MostofthembelongtoOribatidabutsparsein-dividualsofCladoceraandinsectswerealsoidentied.Theyrepresenttherstpre-Quaternaryinvertebratefossilsfoundinsedimentsofcontinentaltemperateclimate.ThePliocene/Pleistoceneageofthesedimentswasdeterminedbypaleomagneticdatingchronologicallycalibratedbymicro-mammalbiostratigraphy.Invertebratefossilscouldbevali-datedasnewproxyforthestudyofcavesedimentsduetotheirsuitabilityforecologicalandpaleogeographiccorrela-tionsincavesandoutsidethecaves.Theyalsobringaddi-tionalinformationaboutcaveformationandkarsthydraulicregimeinthearea.Althoughthenumberofremainswasverylow,itisevidencethatclimaticconditionsincavesal-lowabetterpreservationoffossilremainsofsomegroupsascomparedtomostofthesurfacehabitats.Thismayopenanewdirectioninthestudyofcavesediments. 1IntroductionCavesedimentspreservethegeologicalandpaleoenviron-mentalpastHoracekandBosak,1989aswellasbio-logicalandanthropologicalinformatione.g.,KuklaandLozek,1958;HoracekandLozek,1988;Bosaketal.,1989; Correspondenceto:O.T.Moldovanoanamol@hasdeu.ubbcluj.ro SasowskyandMylroie,2004.Thisisofspecialimpor-tancefortheterrestrialcontinentalhistory,wherecorrel-ativesedimentsaremostlymissingHoracekandBosak,1989,whichisthecaseofthestudiedkarstregion.Cavesedimentsareformedinplaceincaves,orareallochthonousinoriginKyrle,1923;KuklaandLozek,1958.Twocon-trastingfaciescanbedistinguishedamongcaveenviron-mentsKuklaandLozek,1958.Theentrancefaciesin-cludesne-grainedsedimentstransportedfromthevicinityofthecavebywind,waterandslopeprocesses.Itrepre-sentsthemostvaluablesectionofthecavefromastrati-graphicpointofviewasitmaycontaindatablearcheolog-icalandpaleontologicalremainsthatareprotectedfromsur-faceerosion,weatheringandbiochemicalalterationcf.FordandWilliams,1989,2007.Theinteriorfaciesdevelopsinthosepartsofthecavethataremoreremotefromthesur-face.Adominantpartofsedimentsbelongingtotheinteriorfaciesforminvadoseconditions.Sedimentarysequenceshereareextensive,consistingofuvialgravelsandsandsoverlainbyoodorinjectadepositsoflaminatedsiltsandclaysoftenintercalatedbyspeleothems.Theymayalsocon-taindejecta,colluvialmaterialandouterclasticsedimentsincludingmarineones,oftenre-depositedand/orinjectedforlongerdistanceswithinthecavecf.FordandWilliams,1989,2007.Mostofthesedimentsresultfrommaterialcar-riedbysinkingstreamsandgravity-driveninltrationfromthesurfacee.g.,FordandWilliams,1989;BrinkmanandReeder,1995;White,2007.Duetothedynamicen-vironmentofcaveinteriorsandperiodicityofevents,sed-imentarysequencesoftenrepresentaseriesofdepositionalanderosionaleventssedimentarycycles.Thesearesepa-ratedbyunconformitiesbreaksindeposition,whichmay PublishedbyCopernicusPublicationsonbehalfoftheEuropeanGeosciencesUnion.

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1826O.T.Moldovanetal.:Invertebratefossilsfromcavesediments representperiodsofsubstantialduration,andtheerosionalphasesmaybeofmuchlongerdurationthatthedepositionaleventse.g.,Bosak,2002,2008;Bosaketal.,2003.Suchsedimentsreectpastuvialorlacustrineconditionswithinthecave.Distinctsequencesofclasticsedimentsmaybecausedbychangesinwatervolumeandvelocity,sedimentsource,surfaceweatheringconditionsanddepositionalcon-ditionsthatareafunctionofclimatechange.Moreover,spe-cictopoclimaticfeaturesofthecaveenvironmentsdelaythedestructionprocessesoffossilremainsSasowskyandMyl-roie,2004;Polketal.,2007.Therefore,manytypesofin-formationcanbewellpreservedintheunderground,suchas:theabove-groundsourcewhichmaynolongerbefoundonthesurface,theenvironmentalevolutiononthesurface,includinginformationonpastvegetationandlandusewhichisnotpreservedinfast-changingsurfacesoilsKuklaandLozek,1958;Bottrell,1996;CourtyandVal-lverdu,2001;Pannoetal.,2004;Polketal.,2007,thesub-surfacedepositionalprocessesKuklaandLozek,1958,andthelong-lastinggeneralevolutionofthesub-surfaceandsurfacekarstcf.ZupanHajnaetal.,2008b.Inkarstareas,theevolutionofthesurfacelandscapeisrecordedinsidethecaves;loweringofthebaselevelshiftsac-tivespeleogenesistolowerelevations,andtheoverlyingpas-sagesaresubsequentlyabandonedandlledbysediments.Clasticsediments,sometimesintercalatedwithchemicalpre-cipitates,transportedfromthesurfacethroughthecavesarefrequentlypreservedunalteredformillionsofyears,provid-ingdifferenttypesofinformationBosaketal.,1998,2003;Sasowsky,2007.Althoughtherecognitionthatcavesed-imentscanreectandconservepaleoclimaticdataisoldKuklaandLozek,1958andliteratureherein,thescien-ticinterestinpaleoclimaticrecordfromclasticcavede-positsdevelopedonlyinthepastdecadesseesummaryofWhite,2007.Mostlyfossilvertebrateremainshavebeenidentiedinclasticcavesedimentsandusedtocharacter-izeenvironmentalconditions,orusedaspaleoclimateprox-ies.Speleothemshavebeenbroadlyutilizedaspaleoclimaticproxiesusingstableisotopesincombinationwithnumeri-caldating,palynologyandsedimentologicalfeaturesofclas-ticcavesedimentsBastin,1978;Bastinetal.,1986amongothers.Environmentalstudiesofcavesediments,especiallythoseoftheentrancefacies,havebeenalsocarriedoutusingdifferentkindsofproxies;e.g.,charcoal,sedimentpollenandcoprolitepollenCarrionetal.,1992a,b,1995,1997,2005;Finlaysonetal.,2008,stablecarbonisotopesoforganicmat-terTurneyetal.,2001,pesticidecontentderivedfromagri-culturalpollutionBottrell,1996,thepresenceofacertainvegetationtypePannoetal.,2004,thefulvicacidfractionoftheorganicmatterPolketal.,2007,magneticmineralpropertiese.g.,Elwoodetal.,1996;Sroubeketal.,2001,ormollusksincavearcheologicalsites,etc.Otherstudiesthatfocusedonrecentcavesedimentsasaproxyfortheen-vironmentalchangesonthesurfacewerespecicallylinkedtoland-usePolketal.,2007.OnlyasinglepaperPolyaketal.,2001mentionedthediscoveryoftwelvespeciesofori-batidmitesintwoHolocenestalagmitesfromNewMexico,whichwereintegratedinapaleoclimaticstudy.Ourstudyrepresentstherstattempttoidentifyandstudyfossilinvertebratesinclasticsedimentsfromtheinteriorcavefaciesandtodiscusstheirpossibleuseasbiologicalproxiesinpaleoenvironmentalstudies.Similarmethodsandproxiesappliedtoothersedimentarydeposits,suchaslakes,seasandloticenvironments,canalsobeappliedtocavesedimentsinordertoobtaininformationonpaleoclimateandpaleoenvi-ronmentalconditionsatthetimeofsedimentdeposition.Sur-facesedimentarydeposits,especiallylacustrineones,repre-sentpaleoecologicalarchivesofplantmacrofossils,pollen,algaeandfossilinvertebrates.Paleolimnologyhasdevel-opedasamultidisciplinaryscienceespeciallyinthelasttwodecadesusingphysical,chemicalandbiologicalproxiespre-servedinlakesedimentsLuoto,2009.Thestructureandthecompositionoffossilassemblageswepreferthetermfossil,asproposedbyEricksonandPlatt,2007,to“subfossil”varyinresponsetochangesintheenvironment,reectingpastclimate,nutrientconditions,oxygencontent,pH,pollutionorecologicalinteractionsLuoto,2009.ThecladoceransCrustacea;Rautio,2007,chironomidsDiptera;Walker,2001,andostracodsCrustacea;Holmes,2001arethemostcommonlyusedinvertebrateremainsinpaleolimnology.Itisnotdifculttoidentifythemtoaspecieslevel,andtheirau-toecologyiswellknownLuoto,2009.Otherremains,suchasprotozoans,bryozoans,oribatidmitesAcarina,insectsandmollusks,areratherrarelyusedSmol,2002.Untilre-cently,invertebratefossilshavemostlybeenstudiedinlakesedimentsandonlyfewoftheminuviatileenvironmentsGandouinetal.,2006;Engelsetal.,2008;Howardetal.,2009.Clasticsedimentarysequencesbelongingtotheinteriorcavefacieswerecarefullyselectedforarstattempttondfossilinvertebratesincaves.Sedimentarysectionswelldatedbymagnetostratigraphyandpaleomagneticdateswerecali-bratedbybiostratigraphyHoraceketal.,2007;ZupanHajnaetal.,2008b,2010.TheselectedcavesarelocatedintheClassicalKarst,thepartoftheSloveneDinaricregionofKrassoutheasternEurope.ShallowmarineDinaricCar-bonatePlatformdepositsJurassictoPaleogenearecoveredbyEoceneyschsiliciclastics.BothunitsareoverthrustintectonicallycomplicatedstructuresPlacer,1999.Nopost-Eocenemarineorterrestrialdepositsarepreservedonthesurfacenow.Surfacemorphologyandkarstevolvedduringasinglepost-Eocenekarstperiod.Speleogenesiswaslaterfollowedbycaveinllingprocessespartialorcompletefos-silizationthatstartedattheOligocene/MioceneboundaryasindicatedbyssiontrackAFTAandpaleontologically-calibratedpaleomagneticdatafromsomeofthestudiedsitesZupanHajnaetal.,2010. Biogeosciences,8,1825– 1837 ,2011www.biogeosciences.net/8/1825/2011/

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O.T.Moldovanetal.:Invertebratefossilsfromcavesediments1827 Fig.1.LocationofthestudiedsitesinSloveniaseethegeologicalmapintheupperrightcorner;mapafterPlenicaretal.,1969:C=fossilcaveintheCrnoticeQuarrythesectionisrepresentedbyanunroofedcavelledwithyellowuvialsedimentcoveredbyredclaywithowstone,T=TrhlovcaCavetheoldestpartofthesectionwithuvialsediments,R=RaciskapecinaCavethesectionisanalternationofowstoneandclaydeposits;PhotosbyA.Mihevc. 2Materialsandmethods2.1ThestudiedsitesThreereliccaves,alllocatedintheDinaricClassicalKarst,wereselectedforthisstudy–Trhlovca,Raciskapecina,andacaveinCrnoticeQuarryFig.1–alsoduetothefactthatdetailedpaleomagneticdatingwascalibratedbymicromam-malbiostratigraphyintwoofthemHoraceketal.,2007.TrhlovcaCave40018.800N;135604500EbelongstotheDivaskaCaveSystem.Thecaveispartofanancientandmoreextensivesystemcompletelychokedbysediments.ThecavewaslaterpartlyrejuvenatedandsedimentsexhumedasaconsequenceoftheevolutionoftheunderlyingDivaskaCaveBosaketal.,1998,2000.Thepreserveduvialsedimentsandspeleothems,depositedinvadoseconditions,arelocatedinstratigraphicallyrelevantpositionFig.2b.Averticalsec-tion,4.5minthickness,wasdescribedindetailbyZupanHa-jnaetal.b.Itscentralpartiswellstratiedandstartsfromthetopwithabrownish-red,clayeysandwithintercala-tionsoflight-greyishandyellowish-brownsands.Therestofthesectionisrepresentedbymulti-coloredclaysintheupperhalfandchocolate-brownclaysbelow.Inthebasalinterval,siltytoveryne-grainedsandyadmixturesoccurinbandsandlaminae.ThearrangementofRreverseandNnor-malpolarizedmagnetozonesshowsagesolderthan1.77MaZupanHajnaetal.,2008b. www.biogeosciences.net/8/1825/2011/Biogeosciences,8,1825– 1837 ,2011

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1828O.T.Moldovanetal.:Invertebratefossilsfromcavesediments Table1.AlistofinvertebratefossilsinthesamplesofcavesedimentsinSloveniaonlysampleswithidentiableanimalremainsareshown. Site/SampleShortdescriptionofthesedimentIdentiedtaxaNo.individualsObservation 1Trhlovca1T1redclayMiracarusn.sp.1newspecies 2Trhlovca2T2beigeclayOpiellacf.Rhinoppian.sp.1Dissorhinan.sp.11newspeciesspecimeninpoorcondition 3Trhlovca5T5redclayDaphniasp.Dissorhinan.sp.ZygoribatulafrisiaeTetramoriumsp.Insectalarvae11111newspeciescosmopolitecosmopoliteunidentied 4Raciska4R4redclayOppiellaRhinoppian.sp.2Miracarusn.sp.Suctobelbellasp.?421newspeciesnewspeciesspecimeninpoorcondition 5Crnotice1C1yellowclayOrthocladiinaeAstigmatida?11incompletespecimeninpoorcondition RaciskapecinaCave30'12.500N;149'1.5600Erep-resentsarelicofanoldcavesystem.The2mthickverti-calsectionhasacompositestratigraphicthicknessof6.5mHoraceketal.,2007;ZupanHajnaetal.,2008b;Fig.2c.Itslowerpartconsistsofavaultedstalagmitethatincludesseveralinterbeddedlayersofredclays.Itisoverlainbyathickintervalofredclayswithsomesiltyandsandyinterca-lationsandthincalcitecrustsandfossilsvertebratesandin-vertebrate–Potamon.ThemicromammalswithApodemus,cf.BorsodiabelongtomiddletolateMammalNeogenebio-zone17MN17;ca.1.8.4Ma;Horaceketal.,2007;posi-tionofR1andR2inFig.2.Theclaysarepondedandpartlycoveredbyseveralcollapseboulders.Theupperpartofthesectionconsistsofsubhorizontallylaminated,porousandlight-coloredowstonewithsomeancientrimstonedamsandinterbeddedredclaysandsilts.Lutiticinterbedsbetweentheowstonelayersresultedfromsuccessiveoodingthatdepositedwell-sortedne-grainedallochthonoussediments.ThismayindicateeitheradistantpositionfarfromtheponorofthesurfaceriveroranallogenicstreampassingthroughasystemofsumpsHoraceketal.,2007.ThetopofthesectionisowstonewithintercalationsofbrowncaveloamswithbonefragmentsofUrsusspelaeus.FaunafromtheclaypermittedthearrangementoftheinterpretedmagnetozoneswiththeGeomagneticPolarityTimeScaleGPTS;CandeandKent,1995.TheboundaryofN-andR-polarizedmag-netozonewithintheintervalwithfaunawasidentiedwiththebottomoftheC2nOlduvaisubchron.770.950Ma.ThebasalsedimentscanbecorrelatedwiththelowerpartoftheMatuyamachron.150.581MaandtheGausschron.581.580MaHoraceketal.,2007.QuarryoperationsintheCrnoticeQuarry33'56.300N;1352'47.700EuncoveredmanycavescompletelylledwithsedimentsBosaketal.,1999,2004;Mihevc,2001,2007;Fig.2a.Thestudiedsectionislocatedinthewesternquarrywallandrepresentsarelicofanextensive,sediment-lledpassagewithadiameterofabout10mandaheightofmorethan17m.Itstopislledbyspeleothembrecciawithredclaymatrix.Thisisunderlainbyaninterval,upto4.5thick,oflight-colored,laminatedsiltsandclays,sometimessandy,overlyingtherestofthesectionwithdeeperosionandaslightunconformityinside.Thelowermost7mofthecavellarecomposedofcyclically/rhythmicallyarrangedmulti-coloreduvialsedimentsclaystointraclasticmicroconglomerates.ThisllrestsonsessiletubesoftheserpulidMarifugiacavat-icaonthenortherncavewallMihevc,2000;Mihevcetal.,2001.ThemammalremainswithDeinsdorasp.,Bereme-diassidens,Apodemuscf.atavus,Rhagapodemuscf.fre-quens,Glirulussp.,Cseriasp.thatbelongtoMN15–MN16ca.3.0.1Mawerefoundinthesamehorizonastheser-pulidsHoraceketal.,2007.Thebasal1miscomposedofmulti-coloredlaminatedsiltsandclaysdevelopedintwosequencesseparatedbyanangularunconformity.Thefaunaindicatedanageofthellolderthan1.77MabaseoftheC2nOlduvaisubchron.Mostprobably,thellbelongstotheGausschron.6.6MaortheotherN-polarizedsub-chronwithintheGilbertchron4.18.29or4.48.62Ma;Bosaketal.,2004;Horaceketal.,2007. Biogeosciences,8,1825– 1837 ,2011www.biogeosciences.net/8/1825/2011/

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O.T.Moldovanetal.:Invertebratefossilsfromcavesediments1829 Fig.2.Lithologicalsectionsatthestudiedsites:aasectioninthelledhorizontalcavepassagecutbytheCrnoticeQuarrytheoriginalsection:1=limestone,2=owstonemixedwithred-dishclay,3=largeowstoneboulders,4=allogeniclaminatedu-vialsediment,5=wallandsedimentwithtubesofMarifugiaca-vatica,6=samplingpointsC1–C2;modiedfromBosaketal.,2004.bAsectionexposedintheTrhlovcaCave,severalme-tersinthickness:1=limestone,2=reddishyoungerllofclay,siltandnesand,3=clayslighterincolor,withahighersandproportion,3=yellowishbrownsandyclay,4=brownishtoochresandyclay,5=samplingpointsT1–T5;modiedfromBosaketal.,2006;cRaciskapecinaCavesection:1=limestone2=owstone,3=redclay,siltandsand,4=brownclayswithgravelandcavebearbones,5=stalagmitesinowstone,6=speleothemsonthesectionsurface,7=collapsedlimestoneblocks,8=samplingpointsR1–R4;modiedfromZupanHajnaetal.,2008b. 2.2SamplingprotocolsandinvertebratefossilidenticationAtotalof11samplesweretakenfromtheselectedsitesasindicatedinFig.2.Thesamplesweretakenfromtheex-posedfaces,whichhadalreadybeensampledforpaleomag-neticandpaleontologicalstudies.Giventheexploratoryna-tureofourresearch,thesampleswereselectedfromwhatap-pearedtobedistinct,stratigraphicunits,locatedinpositionsthatwouldallowaclearcorrelationwiththepreviouslyes-tablishedpaleontologicalandmagneticchronostratigraphy.IntheCrnoticeQuarry,samplesweretakenonlyfromtheupperpartoftheoriginallydescribedsectionthatremainedintactafterarecentcollapseofthetopmostsedimentsinthequarryface. Fig.3.Lightmicroscopephotographsoftheidentiedfossilin-vertebratesinthecavesedimentsinSlovenia:aDaphniasp.;bOrthocladiinae;cDissorhinan.sp.;dZygoribatulafrisiate;eMiracarusn.sp.;fOpiellacf.Rhinoppian.sp.1;gSucto-belbellasp.?;hOppiellaRhinoppian.sp.2. Approximately1kgofsedimentwastakenfromeachsam-plingpointandplacedinsealedplasticbagswithalabel.Inthelaboratory,thesampleswerekeptin10%KOHfor30min,andwashedsuccessivelythroughsievesof250m,125mand40m.Sub-samplesforeachsievedimensionwereexaminedseparatelyunderanOlympusSZX2stere-omicroscopein90alcoholandeachspecimenwasidenti-edunderanOlympusBX51microscope.Identicationoftheindividualswascarriedoutfollowingthespecicmeth-odsforeachgroup.3ResultsanddiscussionThenumberofidentiableinvertebratefossilsinallsampleswasverylowTable1.Someofthesampleswerecom-pletelyinvertebrate-sterile.Unidentiableanimalfragmentsandvegetalfragmentswerealsofound.Theonlyrelativelywell-preservedspecimensbelongtothegroupsofCladocera www.biogeosciences.net/8/1825/2011/Biogeosciences,8,1825– 1837 ,2011

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1830O.T.Moldovanetal.:Invertebratefossilsfromcavesediments Crustacea,OribatidaAcarina,andChironomidaandHy-menopteraInsecta.Allthesegroupsarecommonlyiden-tiedininvertebratefossilassemblagesfromlakesedimentsElias,2007.3.1Faunalinventory3.1.1CrustaceaCladoceraOneDaphniasp.wasidentiedintheTrhlovcaCave.TheidentiedspecimenisinarelativelygoodconditionFig.3a,butcompletelyattenedlaterally.Daphniaareknownaslarge-bodiedpelagicoffshorecladocerans.KorholaandKorholaetal.foundmaximumlakedepthtobethemostimportantfactorexplainingcladocerandistributioninFennoscandianlakes.Jeppesenetal.foundonlypost-abdominalclaws,mandiblesandephippiaofDaphniaspeciesindeepQuaternarylakesediments.3.1.2AcarinaOribatidaMitesofthesuborderOribatidaAcarina,Arachnidaaretypicalsoil-dwellingmicroarthropods,whichcanbealsofoundincaves.Mitesareofhighpotentialvalueasbioindi-catorsoftheecologicalconditionsinterrestrialandaquaticecosystemsLebrunandvanStraalen,1995;Behan-Pelletier,1999;Gulvik,2007;GergocsandHufnagel,2009.Almostallstudiessupporttheideaofagroupwithmanyrepresen-tativeslivinginhumidhabitats.Droughtsusceptibilityex-ertsitseffectsviafoodlimitationindirectlybyadecreaseofmicrobiotainsoilorothersubstratumasaresultoflackofwater.Nevertheless,somespeciesadaptedalsotoxericcon-ditionsorhighvaluesofhumidity.Theseminutearthropodsareusuallypreservedwellenoughinlacustrineoruvialsed-imentsandinsufcientnumberstobeusefulasproxiesininvestigationsofQuaternarypaleoclimate,paleoecologyandstratigraphySolhyandSolhy,2000;Solhy,2001;Polyaketal.,2001.Fivespecieswereidentiedinthestudiedsam-plesandfourofthemarenewtoscience.AnewspeciesbelongingtoMiracarusMicrozetidaeFig.3ewasfoundinthesedimentsoftheTrhlovcaandRaciskacaves.Modernrepresentativesofthegenusareforest-litterinhabitants.OppiellaRhinoppiasp.1andOp-piellaRhinoppiasp.2Fig.3f,h,twospeciesofthefam-ilyOppidaewereidentiedinthesedimentsoftheTrhlovcaandRaciskacaves.SpeciesofthegenusOpiellasensulatocanbeconsideredoneofthemostcommonarthropodgroupsonEarthNortonandPalmer,1991withhighdiversityandabundanceinforestlitter,alsopresentinshrublands,eco-tonezonesandgrasslands.ThenewspeciesofOppiellafromtheRaciskacavemorphologicallyresemblesaspeciesknownfrommoderncaveenvironments.Thedifferenceisinthelengthofsensilla,beingmuchshorterthanintheex-tantform.ThespeciesisthereforeanextinctelementofthecavefaunaofSlovenia.Dissorhinasp.Fig.3cofthesamefamilyOpiidaewasrepresentedintheTrhlovcaCavebytwospecimens,probablybelongingtothesame,newspecies.Somespeciesofthisgenusprefertheborderbe-tweenforestandopenareasSeniczaketal.,2006.TaylorandWoltersmentionedthetoleranceofthisgenustodrought.ZygoribatulafrisiaeOribatulidaeFig.3dfoundintheTrhlovcaCaveisaspecies,foundtodayinmorearidsettingsShepherdetal.,2002.Thespeciesisxero-tolerant,todayknownfromrepeatedlydrying-outmossesandlichens,ofteninarboricolmicrohabitats.Suctobelbellasp.Sucto-belbidaeFig.3goftheTrhlovcaCaveresemblesspeciesoftherecentgenusSuctobelbellainsomecharactersbutdis-playssomespeciccharacters,whichmaydeneadifferentgenus.Speciesofthisfamilyarecommoninlitterandupperorganiclayerofforestsoils,especiallywithalargeamountofdecomposingorganicmatterandwithabundantfungalhy-phae.Someofthelivingspecieshavebeenfoundinrottingwoodandunderbarkofdeadtrees.3.1.3InsectaChironomidaOnerepresentativeofOrthocladiinaeFig.3bwasidenti-edintheCrnoticeQuarrycavell.Theabsenceofmen-tummadetheidenticationofthelowertaxonimpossi-ble.SubfamilyOrthocladiinaeisagroupofchironomidDipterawhoselarvaepreferlotichabitatswithcoldandwell-oxygenatedwatersDimitriadisandCranston,2001;Walker,2007.Representativesofthisfamilyinhabitcoldandrun-ningstreamsorunstablesandybottomsoflakes,butaregen-erallyadaptedtolowfoodWalker,2007andareintoler-anttolowoxygenlevelsEggermontetal.,2008.Thisisawidelydistributedfamilyofchironomidsanditsrepresenta-tivesarefrequentlyfoundinlakesediments.3.1.4InsectaHymenopteraOneindividualofthegenusTetramoriumwasfoundinthesedimentsoftheTrhlovcaCave.Thegenusisatypicalinhab-itantofdrylandscapeswithshrubs.Theindividuallacksthehead,butisotherwisewellpreserved.Itrepresentsagenuswithverylargedistributioninthepresentfauna.3.2PaleoenvironmentalsignicanceofthefossilinvertebratesFivetaxaidentiedarenewtoscienceandtheirdescrip-tionisinprogress.OnlyasingletaxonasyetidentiedataspecieslevelisfoundinthemodernfaunaZygoribatulafrisiae.TwotaxawereidentiedatagenuslevelDaphniaandTetramoriumandonetaxonassubfamilyOrthocladi-inae,duetotheirpoorconservation.ThecaverichestinfossilswasTrhlovcaCavewithsixidentiedtaxainthreeofthevesamples.ThemaximumnumberoftaxabelongstotheoldestsampleT5,presumablyduetoclimaticcondi-tionsandsedimentationprocessesatthetimeofdepositionandsubsequently. Biogeosciences,8,1825– 1837 ,2011www.biogeosciences.net/8/1825/2011/

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O.T.Moldovanetal.:Invertebratefossilsfromcavesediments1831 Fig.4.AcorrelationoftheobtainedmagnetostratigraphicresultswiththestandardpaleomagneticscaleGPTS;afterCandeandKent,1995;modiedafterHoraceketal.,2007,mammalzones,temperatureestimatesafterLisieckiandRaymo,2005andposi-tionofthesampleswithinvertebratesinredandvertebratesinblue:T=TrhlovcaCave,R=RaciskaCave,C=fossilcaveintheCrnoticeQuarry. AlltheanalyzedsamplesweretakenfromcavesedimentsdepositedduringthePlioceneandPleistoceneperiods,asdatedbymagnetostratigraphyandpaleontologicalcontentFig.4.DuringthePliocene,theclimateisgloballyknowntohavebecomecooleranddrierthanduringtheMioceneRobinsonetal.,2008,followedbythemorepronouncedPleistocenecooling.Broadly,twoclimaticstagesaredenedintheperiodcoveredbythedepositionofclasticsedimentsinthesampledcavesFig.5:theEarly–MiddlePlioceneca.5.3Mato3.6Mawithhighertemperatures,andtheLatePliocene–EarlyPleistoceneca.3.6to1.77Mawithlowertemperatures,moresimilartothepresent-dayclimateHay-wood,2009.MostoftheinvertebratefossilswerefoundintheTrhlovcaCave.Samples1belongtoamorerecentperiodthantheolderinllingwheresamples4werecollected,butallthestudiedsamplesareolderthan1.77MaZupanHajnaetal.,2008b,2010.ClaysamplesT1andT2includetaxathataretypicalforforesthabitatswithrelativelyhighamountsofdeadorganicmatterintheupperhorizonofsoilMiracarusandOppiellaortaxawhichcanbefoundintheecotonezonesorinopen,moderatelyhumidareasDissorhina.Fromtheoldestsamples,T4–T5,onlyT5containedfossilre-mains.Theecotonezone–typicalDissorhinawasfoundinassociationwithdryhabitattaxa,suchasZygoribatulaandTetramorium.Fossilsfromthesethreesamplesindicateatransitionfromawarmerperiodwithrelativelydriervegeta-tionEarlyorMiddlePliocenetoamorehumidandforestedhabitatsoftheLatePliocene/EarlyPleistocene.ThepresenceofDaphniasp.insampleT5suggeststhepresenceofalakeoralowhydraulicregimeofthestream,aswellastheprox-imityofcaveentrance.Thisassumptionisalsosupportedbythevarvedsedimentsthatindicatecontinuousdepositioninthepast,duringsingle-oodeventsoroodpulsesthatprob-ablylastedlessthanafewthousandyearsZupanHajnaetal.,2008a.Therelativelygoodpreservationofthisclado-ceranisalsoindicativeofaslowowfromthesurfacedowntoplaceofdeposition,possiblyduetothestillepiphreaticpositionofsomepartsofthesubterraneansystemMihevc,2007andtotheshort-distancetransport.ThepresenceofDaphniaspecies,asanindicatorofoligotrophiclowfoodenvironmentsSzeroczynskaandZawisza,2005,supportsthehypothesisofadriervegetationonthesurface.OnlysampleR4providedfossilinvertebratesinRaciskapecina.Thesearerepresentedbythreeindividualsoforib-atidmitesbelongingtothesameecologicalgroupofforestinhabitants.SampleR4comesfromtheupperhalfofthesectionofthiscavethatbelongstotheEarlyPleistoceneter-minationoftheOlduvai-chron.TheonlychironomididentiedatafamilylevelwasfoundinsampleC1inCrnoticeQuarry.Locatedintheupperhalfofthesedimentsection,theOrthocladiinaedipterancanbeusedasanindicatorforcolderandwell-oxygenatedwaters.Theindividualwascarriedundergroundbyarelativelyfastowingriver,whichsuggestsacolderclimatewithmorein-tenseprecipitationtowardstheendofthePliocene.3.3CorrelationswithotherproxiesThediscoveryofvenewspeciesandperhapsanewgenusisimportantnotonlyforthetaxonomyofthegroupofmitesinthiscase,butisalsoofpaleoenvironmentalsig-nicance.ThePliocene/Pleistocenecavesedimentsshow www.biogeosciences.net/8/1825/2011/Biogeosciences,8,1825– 1837 ,2011

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1832O.T.Moldovanetal.:Invertebratefossilsfromcavesediments Fig.5.Invertebratefossilsfoundinthestudiedcaves,correlatedwithclimate,vegetationandfossilvertebrates. averylowconcentrationoftaxa,ornoneatallinmorethanhalfoftheanalyzedsamples.Thescarcityoffaunaremainsmakesfurtherbiostratigraphicinterpretationimpos-sible,butthepresenceandecologyofsometaxacanbecorrelatedwiththediscoveredvertebrateremainsaswellaswiththeclimateandcorrespondingvegetationofthediffer-entPliocene/Pleistoceneperiods,indicatingatransitionfromadryormildclimatewithmixedforesttoacolderclimateandmorehumidforestTable2,Fig.5.Theuseoflessfre-quentlyoccurringzoologicalremainsinpaleolimnologywasemphasizedbyLuotoasanimportantproxyforenvi-ronmentalchanges.ThefossilvertebrateassemblagefromtheCrnoticeQuarry,includingDeinsdorasp.,Beremendiassidens,Apodemuscf.atavus,Rhagapodemuscf.frequens,Gliru-lussp.andCseriasp.Horaceketal.,2007,istypicalforatemperateandhumidclimate.Thehighnumberoffor-estandshrubinhabitantsApodemus,Glirulus,Myodesgen-era,togetherwithtaxathatarewidelydistributedintemper-ateandhumidclimatesBeremendiassidens,BlarinoidesandDeinsdoragenera,isindicativeofamildclimate.AtRaciskapecinaCave,themostimportantdifferenceisthepresenceofMyodesandBorsodiaHoraceketal.,2007asindicatorsofacolderanddrierclimate.Owingtothescarcepresenceofidentiableinvertebrates,acorrelationwiththeequallyscarcevertebraterecordisproblematic.OnlythreeoftheinvertebratesamplesC1,T1andR4comefromthesameperiodsasthefoundvertebratefossils,andcor-relationsweremadeonlyforthesesamples.SampleC1providesasingleindicatorforcold/well-oxygenatedwater Biogeosciences,8,1825– 1837 ,2011www.biogeosciences.net/8/1825/2011/

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O.T.Moldovanetal.:Invertebratefossilsfromcavesediments1833 Table2.TaxafoundincavesedimentsofSloveniawiththecorrespondingvegetation,sedimenttypeandorigin,andvertebratefossils. StageTAXAEnvironmentSedimentVertebrates TrhlovcaRaciskaCrnoticeorigin UpperPliocene/LowerPleistoceneMiracarusOpiellaDissorhina––forest,ecotonezonessurface,uvial––OppiellaMiracarusSuctobelbellaPotamon–forest,ecotonezones,grasslands+riversurface,uvialshteeth,micromammalslivinginsteppeandtreehabitats––OrthocladiinaeMarifugiacavaticacold,welloxygenatedwatersurface,uvialshteeth,micromammalslivingintreeandriversidehabitats LowerPlioceneDaphnia,Dissorhina,ZygoribatulaTetramolium––drylandscapewithshrubs+riversurface,uvial– MentionedinHoraceketal.. Orthocladiinae,whichissupportedbythepresenceofBere-mendia,Rhagapodemustypicalfortemperateclimate,andofCseriaandMimomysthatarewater-relatedmicro-mammals.SamplesT1andR4includeOribatidainvertebratesgeneraOppiella,MiracarusandSuctobelbellathatinhabitforestlitter,shrublands,ecotonezonesandgrasslands,andverte-brateswhichindicatethepresenceoftreesinwetforestsandsteppichabitats,typicalhabitatsforMyodesandBorsodia.Thepresenceofinvertebratedrought-toleranttaxainT5in-dicatesalandscapewithshrubsandgrasslandduringthelateEarlyPliocene.ThepresenceofforestinhabitantsinT1andR4correlatedwiththepresenceofcold/dry-associatedverte-bratespointstoacolderclimateduringtheMid-Pleistocene.Betweenthesetwochronologicallyextremesamples,inver-tebrateswerealsoidentiedinsamplesC1andT2.Sam-pleC1correspondstotheadventoftheLatePliocenecool-ingandprobablyhigherowratesonthesurface,whichex-plainsthepresenceofthechironomid.SampleT2tswithinthepictureofforestsmixedwithopenareasduringtheEarlyPleistocene.Nopollenwasfoundinthestudiedsamplesandsec-tions,butdetailedstudiesonthenearbyItalianandperi-MediterraneansitesgiveninBertinicanbewellcor-relatedwiththeinvertebratefossilsfoundintheSloveniancavesedimentsTable2.DuringtheZanclean,thecli-matewaswarmeranddrierormorehumidthantoday,de-pendingonthegeographicallatitudeandaltitudeFaque-tteetal.,2006;foresttaxaofhumidsubtropicaltowarmtemperatezonesweredominantBertini,2010.Thegen-eraldecreaseofhumidityandinstallationofdrierconditionsduringthePiacenzianoccurredpriorto2.5MaBertoldietal.,1989.Alternationsofforestvegetationandsteppes,withtypicalopenvegetationphasescharacterizedthefol-lowingearlyPleistocene,duringtheGelasianandtheCal-abrianBertini,2010.Ingeneral,theoralcompositionandstructureofvegetationofthePliocene-PleistocenearecharacterizedbytherareappearanceofnewspeciesandbyspatiallyasynchronouseventsofdisappearanceofthesametaxonBertini,2010.Frequentchangesinbothclimateandvegetationcanexplaintheextinctionofsomeoftheidenti-edinvertebratetaxainthecavesedimentsofSlovenia.Ingeneral,faunalchangesdependontheenvironment,denedmostlybyclimateandvegetalassociations.Ifchangesarefrequentandofrelativelyshortduration,someofthetaxathatareunabletoadaptmaygetextinct,orevolveintodifferentspecies.Oribatidaisagroupwithalowermobilitythany-inginvertebratespecies,andtheirmigrationcanonlybeovershortdistances,followingtheslowlymigratingvegetationorvegetalassociations.TheidentiednewOribatidaspeciescanbeconsideredeithermembersofanextinctfaunalgroupwithnorelativesinthepresentfaunaofsouthernSlovenia,orancestorsofalivingspecieswhichhaveundergoneevolutioninthepresent-dayhabitatsorindifferentareasnearby.Veg-etationsuccessionintimecontributedtothesettlementofacomplexmosaicofdifferentclimates/biomesintheMediter-raneanzoneBertini,2010.Thissituationdocumentedfororaissupportedbytheinvertebratesfoundinthecavesed-iments.Thesectionthatprovidedmostofthespecimens, www.biogeosciences.net/8/1825/2011/Biogeosciences,8,1825– 1837 ,2011

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1834O.T.Moldovanetal.:Invertebratefossilsfromcavesediments intheTrhlovcaCave,yieldedtaxaindicativeofmixturesofforestswithopendryareas.AsalreadydocumentedbyFin-laysonetal.formorerecentsediments,environmen-talchangesmustbetakenintoaccountatsmallormediumscales.Invertebratefossilscanprovideinformationatsmallscalesandmustbeassociatedwithotherproxiesandprefer-ablywithmoresitesofthesameregioninordertoprovidepaleoenvironmentalinformationthatwouldbesignicantatregionalscale.3.4ImplicationsforthereconstructionofkarstevolutionIfthestandingconditionofthefossilisknown,itsendpointcanbeusedtoinfersomeofthegeologicalprocessesthatactedonthebodyduringitstransporttothesiteofdeposi-tionErickson,1988.Thepresenceofsurfaceanimalsinpre-QuaternaryinteriorfaciesofcavesedimentsisindicativeforanintensekarstevolutionoftheDinarickarst,includ-ingthellingofapartofthecavitieswithsedimentduringtheNeogene.Theverylowabundanceoffossilsdepositedinclasticcavesedimentsoftheinteriorfaciesiswellknowne.g.,HoracekandLozek,1988.Thisisprobablyrelatedtotherunoffinthedrainageareaortothehydraulicsofthesubterraneanstreams.Almostallofthestudiedinver-tebratesampleshaveatleastoneaquaticcomponent,andallwithasinglepossibleexceptionaresurfaceinvertebrates,whichwerebroughtinsidethecavesbyallogenicstreamsanddepositedtogetherwiththesediments.Themostcom-monclasticsedimentsinthestudiedcavesaredifferenttypesofnelaminatedclaysandsilts.TheyweredepositedfromoodwatersuspendedloadunderconditionsofpulsedoworcavelakescomparablewithslackwaterfaciesofBoschandWhite.Thisdepositionalprocesscorrespondstomoreorlessregularoodingofkarstareasbysinkingrivers.MineralassemblagesofthecavedepositswerederivedfromhighlyhomogenizedweatheringproductsofEoceneyschsedimentsandsoilcoveronPaleoceneandCretaceouslime-stonesZupanHajnaetal.,2008b.Thetectonically-drivenloweringoftheregionalbaselevelwasconnectedwiththechangeintectonicregimeatca.6MaVrabecandFodor,2006.Thiscausedatransitionfromanepiphreatictoava-doseregime,followedbyadecreaseinhydraulicheadinsomecavesystemsortheirhigher-situatedpartsandacom-pletellwithuvialsedimentsasaconsequenceMihevc,2007;ZupanHajnaetal.,2008b.Asalreadymentioned,thepresenceofDaphniasp.anditsrelativelygoodpreservationinsampleT5indicatestheexistenceofasystemwithlowhydraulicheadatthecaveen-tranceandmultipleoodeventsthatcontributedtothecon-tinuousdepositionofvarvedand/orcyclicallyarrangedsed-iments.ThechironomidinsampleC1indicatesaperiodofrelativelyhighowrateinavadoseregime.ThechironomidwasfoundatthesamelevelasMarifugiacavatica,serpulidpolychaetesattachedtowallsattheair/karst-waterinterface,stilllivinginthecavesoftheDinarickarstforsummaryseeMihevcetal.,2001.Pre-Quaternaryarthropodremains,abundantinmarinesettingsandquitecommoninamber,arerareandhaveyetbeenreportedincontinentalsedimentsonlyfromArcticen-vironmentsseeacompletelistinElias,2010.Thedeposi-tionalmechanismsandthelowintensityofbiochemicalpro-cessescanexplaintherelativelygoodstateofpreservationofoldinvertebrateremainsbothinthecavesedimentsandArcticlakesediments.Althoughthenumberofidentiedin-vertebratesatthestudiedSloveniansitesissmall,theirstateofpreservationisrelativelygoodconsideringtheageofthesediments.Thismaysuggestsacombinationof:arela-tivelyshortandslowtransporttothesiteofdeposition,2arapidburial,i.e.,ahighsedimentationrate,andsubduedmicrobialandbiochemicalprocessesthatcouldhavealteredtheentireorganisms.4ConclusionsThreesitesofreliccavesfromtheClassicalKarstofSloveniaproviderstevidenceofPliocene/Pleistoceneinvertebrateremainsincontinentalclasticsedimentsfrominteriorcavefaciesintemperateregions.Thesendssuggestthatcavesedimentscanpreserveyetanotherproxyfortheassessmentofpaleoclimaticandpaleoenvironmentalconditions.Thisnewproxymaybeimportantforanumberofreasons:ithescarcityofvertebratefossilsinpre-Quaternarycavedeposits;iitheneedforcross-validationofinferredpaleoenviron-mentalsettingsinthecaseofincompleteorambiguousco-evalproxies;iiithecaveconditionsthatarepronetoabetterpreservationofoldchitinousinvertebrateswhencomparedtosurfacesettings;ivitsadded-valueasasourceofin-formationonspeleogenesisandevolutionofkarsthydraulicregimesevenforpaleokarstsettings.Cavesareknownassystemswithlow-energyinputfromthesurfaceandlow-energyinsituproduction.Thismayhampertheuseoffossilinvertebratesfromcavesduetotherelativelylowchanceofndingidentiablecaveorsurfacespecimenstransportedintocaves.However,forthethreedescribedsites,arelativelylownumberofin-vertebratespecimensweresuccessfullyidentiedanddatedbybiostratigraphy-calibratedmagnetostratigraphybacktoca.1.77toca.4.8Ma,i.e.,acrossthePliocene/Pleistoceneboundary.Whencross-correlatedwiththecoevalvertebratefauna,thesendingssupporttheideaofacolderphaseas-sociatedwiththePliocene/Pleistocenetransition,whilealsobringingnewinsightsontheregionalkarstevolution. Acknowledgements. TheauthorsaregratefultoIlseBartsch,KarinaBattes,SteveBrook,MirelaCmpean,BalintMarko,LauraMomeuforthehelpwithtaxarecognitionandecologicalinformation.JirAdamovichaskindlyhelpedwiththeEnglishrevisionofthemanuscript.Thisstudywasfundedthroughthe Biogeosciences,8,1825– 1837 ,2011www.biogeosciences.net/8/1825/2011/

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O.T.Moldovanetal.:Invertebratefossilsfromcavesediments1835 KARSTHIVESProjectPCCE ID 31/2010fundedbyCNCSIS-UEFISCDIRomania;grantedtoSC,GrantAgencyASCRNo.IAA300130701Paleomagneticresearchofkarstsediments:paleotectonicandgeomorphologicalimplications;grantedtoPBandtheInstitutionalResearchPlanNo.AV0Z30130516oftheGLIASCR,v.v.i.TheauthorsthanktoeditorWolfgangKiesslingandreviewersIraSasowskiandJohnHolsingerfortheirinsightfulcomments.Editedby:W.KiesslingReferences Bastin,B.:L'analysepolliniquedesstalagmites:unenouvellepos-sibilited'approchedesuctuationsclimatiquesduQuaternaire,Annals.Soc.Geol.Belgique,102,13–19,1978. Bastin,B.,Cordy,J.-M.,Gewelt,M.,andOtte,M.:Fluctuationscli-matiqueenregistreesdepui250000ansdanslecouchesderem-plissagedelaGrotteScladinaProvincedeNamur,Belgique,Bull.Assoc.Franc.EtudeQuatern.,1/2,168,1986. Behan-Pelletier,V.M.:Oribatidmitebiodiversityinagroecosys-tems:roleforBioindication,Agr.Ecosyst.Environ.,74,411–423,1999. Bertini,A.:PliocenetoPleistocenepalynooraandvegetationinItaly:Stateoftheart,Quatern.Int.,225,5–24,2010. Bertoldi,R.,Rio,D.,andThunell,R.:Pliocene-Pleistocenevegeta-tionalandclimaticevolutionofthesouth-centralMediterranean,Palaeogeogr.Palaeocl.,72,263,1989. Bosak,P.:Karstprocessesfromthebeginningtotheend:howcantheybedated?,in:EvolutionofKarst:FromPrekarsttoCessation,editedby:Gabrovsek,F.,Carsologica,ZalozbaZRC,Postojna-Ljubljana,2002. Bosak,P.:Karstprocessesandtime,Geologos,14,15,2008. Bosak,P.,Ford,D.C.,Gazek,J.,andHoracek,I.Eds.:Pale-okarst.ASystematicandRegionalReview,Elsevier-Academia,Amsterdam-Praha,1989. Bosak,P.,Pruner,P.,andZupanHajna,N.:PaleomagneticresearchofcavesedimentsinSWSlovenia,ActaCarsologica,27,151–179,1998. Bosak,P.,Mihevc,A.,Pruner,P.,Melka,K.,Venhodova,D.,andLangrova,A.:CavellintheCrnoticeQuarry,SWSlovenia:Palaeomagnetic,mineralogicalandgeochemicalstudy,ActaCar-sologica,28,15,1999. Bosak,P.,Pruner,P.,Mihevc,A.,andZupanHajna,N.:Magne-tostratigraphyandunconformitiesincavesediments:casestudyfromtheClassicalKarst,SWSlovenia,Geologos,5,13,2000. Bosak,P.,Pruner,P.,andKadlec,J.:Magnetostratigraphyofcavesediments:applicationandlimits,Stud.Geophys.Geod.,47,301,2003. Bosak,P.,Mihevc,A.,andPruner,P.:GeomorphologicalevolutionofthePodgorskiKarst,SWSlovenia:Contributionofmagne-tostratigraphicresearchoftheCrnoticeIIsitewithMarifugiasp.,ActaCarsologica,33,175,2004. Bosch,R.F.andWhite,W.B.:Lithofaciesandtransportofclas-ticsedimentsinkarsticaquifers,in:StudiesofCaveSedi-ments.PhysicalandChemicalRecordsofPaleoclimate,editedby:Sasowsky,I.D.andMylroie,J.,KluwerAcademic/PlenumPublishers,NewYork,2004. Bottrell,S.:Organiccarbonconcentrationprolesinrecentcavesediments:Recordsofagriculturalpollutionordiagenesis?,En-viron.Pollut.,91,325,1996. Brinkman,R.andReeder,P.:Therelationshipbetweensurfacesoilsandcavesediments:anexamplefromwestcentralFlorida,USA,CaveKarstSci.,22,95,1995. 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