DoesthemiddleEoceneRíoPichileufú florafromPatagonia,Argentina record initialfloristicresponsetoglobalcoolingandSouthAmericanIsolation? ClaireCleveland DepartmentofGeosciences,PennsylvaniaStateUniversity,UniversityPark,PA16802 [email protected] Introduction SampleofLeafDiversity SouthernHemispherefossilplantsindicatearainforestbiomeacrossmuchofLateCretaceous andEarlyPaleogeneGondwana,butglobalcoolingandseparationofSouthAmericaand AustraliafromAntarcticaarelinkedtosignificantecologicalchangeinthemiddleEocene throughtheOligocene.TheRíoPichileufú fossilflora(RP)fromRíoNegroProvince,Argentina (Fig.1)offersanexceptionalopportunitytoobservetheearliestsignalsofecologicalchange inmiddleEocenePatagonia.TheprincipalrecordofPatagonianfossilplantsduringtheEarly EoceneClimaticOptimum(EECO)istheLagunadelHunco fossilflora(LH)inChubutProvince, Argentinadated52.22(±0.22)Ma(Wilf,2012).RPisdatedca.3MaaftertheEECOat47.74 ±0.05Ma(Wilf2012),duringtheveryfirststagesofcoolingandseparationofSouthAmerica fromAntarctica(Fig2).Althoughthe1938E.W.BerryRPmonographstillrepresentsthemost diversedescribedCenozoicfossilflorafromSouthAmerica,itprovidesonlyaqualitative accountoftheassemblageandjustafractionofitsspecimenshavereceivedmodern paleobotanical treatment.AcomprehensiveupdateoftheRPassemblagecombinedwith quantitativedatafromrecentunbiasedcollectionswillprovidethedataneededtocompareRP withLHandtestforsignalsoffloralresponsetocoolinginPatagoniaandSouthAmerican biogeographicisolation. Objectives Toprepareforquantitativetestsofpaleofloral responsetocoolingandbiogeographicchange,I beginwithanupdateoftheRPflora.RPdiversitywillbeestimatedusingmorphotypes and systematiccomparisonswithtypematerialtoimproveourunderstandingoffloralcomposition. Figure5.Asamplingoffloral diversityandpreliminary identificationsfromtheRío Pichileufú collections (Preliminaryidentification, Morphotype No.,SpecimenNo.). A. Retrophyllum sp.,RI097, BAR_4705 B. Unidentified,RI086, BAR_4595 C. Unidentified,RI039, RP3_0827 D. Unidentified,RI073, BAR_4639 E. Cochlospermum previtifolium Berry,RI071,RP3_0942 F. Unidentified,RI071, RP3_0942 G. “Banera”prehernandiensis Berry, RI082,BAR_4622 H. “Villaresia”congonhifolia Berry,RI080,RP3_1111 I. “Nectandra”prolifica Berry, RI048,BAR_4450 J. Myrtaceae,RI012,BAR_4356 K. Myrtaceae with infructescences attached, RI087,BAR_4706 L. Fabaceae,RI052,RP3_0739 B • 400+RPspecimensfromtheEWBerry1938TypeandCohort collectionattheSmithsonianNationalMuseumofNatural History Figure1.RíoPichileufú andLagunadelHunco locationsmappedonto50 Mapositionsofmodern coastline.Figureadapted fromWilfetal.,2005. C D Río Pichileufú (47.74±0.05Ma) Laguna del Hunco (52.22±0.22Ma) • Allcollectionsquarriedfromtuffaceous caldera-lake sedimentsofHuitrera Formationindicatingsimilar paleoenvironments RíoPichileufú LeafDatabase E Figure2.Siteagesplacedon globaldeep-seaoxygenisotope recordsadaptedfromZachos et al.,2001. Specimens F H I LeafArchitecture F Berry(1938)originallyassignedtheRPfloratoaMioceneageand highlighteditsaffinitiestobothextantSouthAmericanandAustralasian floras.Incontrast,recent40Ar/39Ardatesshowthatthefloraisearliest middleEocene(Wilf,2012),andsystematicrevisionsofangiospermsand conifers(Fig.4)havegreatlystrengthenedAustralasianlinks.Severalof theconifertaxa,suchasPapuacedrus andDacrycarpus,provideevidence supportingahumidandeverwet environmentwithlinkstoextantWest Pacificmontaneforests(Wilfetal.,2009;Wilf,2012).Theflowerofbasal Asteraceae Raiguenrayun cura fromanotherRíoPichileufú locality (Barreda etal.,2010;Barreda etal.,2012)representsa paleobiogeographic linkbetweenSouthAmericaandAfrica(Bandoni De Oliveiraetal.,2009).Insightsfromawell-developedunderstandingofRP anditscomparisonwithLHwillbeusedtoinvestigatetransitionfrom AncientGondwanantoextant-SouthAmericanaffinitiesandtestthe proposedpaleogeographic explanations. Figure4.SomeRíoPichileufú taxathathavebeenrevisedsinceBerry’s1938monograph.A.Acmopyle engelhardti (Berry)Florin1940; specimenRP3_1068.B.Papuacedrus prechilensis (Berry)Wilfetal.2009;specimenBAR-4742a. C.Dacrycarpus puertae Wilf2012;USNM. 40381. D.Agathis zamunerae Wilf2014;USNM-40378.E. Atherospermophyllum guinazui (Berry)CLKnight2013;USNM-40403. F.Ginkgoites patagonicus Villar deSeoane etal.2015;specimenRP3_1112. . PreviousworkindicatesthatalthoughmanyplanttaxaarefoundatbothRPandLH,including gymnospermssuchasDacrycarpus andAgathis,RPlackstaxaabundantatLH,including angiospermsEucalyptus andGymnostoma.Also,taxaobservedatRParenotrepresentedatLH, includingAsteraceae flowers(Barreda etal.,2012).Thisworkwillprovidethefoundationfora comprehensiveupdateofBerry’sRPmonographandquantitativecomparisonoftheRPandLH assemblagesinthecontextofglobalcoolingandSouthAmericanbiogeographicisolation.I predictnetlossofancientGondwananrainforestassociationsandnetgainofcladeswith affinitiestoextantSouthAmericaflorasaswellasshiftsinrelativeabundance. LiteratureCited Figure3.Aprojectdatabase,developedusingFileMakerPro Advancedv.14.0.4,organizesleafdatainarelationalframeworkand generatesautomateddescriptionstoassistmanuscriptpreparation. TerminologyfromtheManualofLeafArchitecture,2009(Ellisetal., 2009)isusedtoensureconsistenttraitcomparisons. L Discussion Images Morphotypes K J G FloralChangeinEocenePatagonia A • 4300+LHspecimensintherecentlypublishedcollectionat theMuseo Paleontológico Egidio Feruglio (Trelew,Argentina) D C E A Materials • 1200+RPspecimensfromtheunbiased2002and2005 collectionsattheMuseo Paleontólogico Bariloche (Bariloche, Argentina) B Bandoni deOliveiraF, CassolaME,Marroig G(2009).In:GarberPA,EstradaA,Bicca-MarquesCJ, Heyman EW,Strier KBeds.NewYork:SpringerScience,55-68. Barreda VD,Palazzesi L,Katinas L,Crisci JV,Telleria MC,etal.(2012).AnnalsofBotany109:127134. BerryEW(1938).GeologicalSocietyofAmericaSpecialPapers12:1-140. EllisB,DalyDC,HickeyLJ,JohnsonKR,MitchellJD,etal.(2009).ManualofLeafArchitecture. CornellUniversityPress,Ithaca,NewYork216pp. FlorinR (1940).Svensk Botanisk Tidskrift 34:117-140. KnightCL andWilfP(2013).Palaeontologia Electronica16:29A-39. Villar deSeoane L,Cúneo NR,Escapa I,WilfP,GandolfoMA(2015).InternationalJournalofPlant Sciences176:346-363. WilfP,JohnsonKR,Cúneo NR,SmithME,SingerBS,GandolfoMA(2005).AmericanNaturalist 165:634-650. WilfP,LittleSA,IglesiasA,DelCarmenZamaloa M,GandolfoMA,etal.(2009).AmericanJournal ofBotany96:2031-2047. WilfP (2012).AmericanJournalofBotany99:562-584. WilfP,Escapa IH,Cúneo NR,Kooyman RM,JohnsonKR,IglesiasA(2014).AmericanJournalof Botany101:156-179. Zachos JC,DickensGR,Zeebe RE(2008).Nature451:279-283. Acknowledgements FundingprovidedbytheNationalScienceFoundation(NSF DEB-1556666andGraduateResearchFellowshipProgram), theEvolvingEarthFoundation,theGeologicalSocietyof America,ThePaleontologicalSociety,andthePaulD.Krynine MemorialFund,PennStateGeosciences.Manythanksfor accesstothecollections,labspace,andabundanttechnical supportprovidedbytheMuseo Paleontólogico Egidio Feruglio (MEF)andMuseo Paleontólogico Bariloche (BAR). MysincereappreciationtoAnnaWhitakerforherassistance attheMEFandforAaronCleveland’sassistancewith photography.Thisworkwouldnotbepossiblewithoutthe supportofmyadvisor,PeterWilf.
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