The rodents from Santpedor-2 (Ebro Basin, NE Spain) confirm the Oligocene age of the latest primates from the Paleogene of Europe

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The rodents from Santpedor-2 (Ebro Basin, NE Spain)

confirm the Oligocene age of the latest primates from

the Paleogene of Europe

Monique Vianey-Liaud, Raef Minwer-Barakat, Laurent Marivaux

To cite this version:

Monique Vianey-Liaud, Raef Minwer-Barakat, Laurent Marivaux. The rodents from Santpedor-2

(Ebro Basin, NE Spain) confirm the Oligocene age of the latest primates from the Paleogene of Europe.

Geobios, Elsevier Masson, 2019, 55, pp.77-88. �10.1016/j.geobios.2019.06.002�. �hal-02263896�

HAL Id: hal-02263896

https://hal.umontpellier.fr/hal-02263896

Submitted on 6 Aug 2019

HAL is a multi-disciplinary open access

archive for the deposit and dissemination of

sci-entific research documents, whether they are

pub-lished or not. The documents may come from

teaching and research institutions in France or

abroad, or from public or private research centers.

L’archive ouverte pluridisciplinaire HAL, est

destinée au dépôt et à la diffusion de documents

scientifiques de niveau recherche, publiés ou non,

émanant des établissements d’enseignement et de

recherche français ou étrangers, des laboratoires

publics ou privés.

The rodents from Santpedor-2 (Ebro Basin, NE Spain)

confirm the Oligocene age of the latest primates from

the Paleogene of Europe

Monique Vianey-Liaud, Raef Minwer-Barakat, Laurent Marivaux

To cite this version:

Monique Vianey-Liaud, Raef Minwer-Barakat, Laurent Marivaux. The rodents from Santpedor-2

(Ebro Basin, NE Spain) confirm the Oligocene age of the latest primates from the Paleogene of

Europe. Geobios, Elsevier Masson, In press, �10.1016/j.geobios.2019.06.002�. �hal-02263896�

Original

article

The

rodents

from

Santpedor-2

(Ebro

Basin,

NE

Spain)

confirm

the

Oligocene

age

of

the

latest

primates

from

the

Paleogene

of

Europe

§

Monique

Vianey-Liaud

,

Raef

Minwer-Barakat

*

,

Laurent

Marivaux

a

InstitutdesSciencesdel’E´volutiondeMontpellier(ISE-M),UMR5554,CNRS,UM,IRD,EPHE,c.c.064,Universite´ deMontpellier,placeEuge`ne-Bataillon, 34095MontpellierCedex05,France

b_{InstitutCatala` dePaleontologiaMiquelCrusafont,UniversitatAuto`nomadeBarcelona,CerdanyoladelValle`s,Barcelona08193,Spain} c_{DepartamentodeEstratigrafı´ayPaleontologı´a,UniversidaddeGranada,Avda.FuenteNuevas/n,18071Granada,Spain}

1. Introduction

ThefossilsiteofSantpedor (EbroBasin,NESpain),originally namedas‘‘Costadela Vila’’,wasfirstmentionedin the1950’s, whenMasachsetal.(1954)brieflydescribedthepresenceoftwo perissodactylsandanartiodactyl:Paleotheriummedium, Plagiolo-phus annectens, and an undetermined Anoplotheriidae (later referredtoAnoplotheriumcommunebyGolpe,1982).Theseauthors assignedthelocalitytothelateEocene.Inthe1980’s,theteamsof the Institut de Paleontologia de Sabadell (now Institut Catala` de

Paleontologia Miquel Crusafont, ICP) and the Universidad de Barcelona decided to resumethe study of this locality. On the onehand,theyrevisitedthemacromammalremainsfirstreported byMasachsetal.(1954),notingthatthematerialwassoscarce thatitcouldnotbedeterminedatthespecificlevelanddidnot allowforaprecisedatingofthelocality.Therefore,accordingto Arbiol and Sa´ez (1988), the faunal list of the classical site of Santpedor consisted of Paleotherium sp., Plagiolophus sp. and Anoplotheriidaeindet.Ontheotherhand,theseteamsreturned andprospectedtheareainordertofindlevelswithmicromammal remains.Agustı´ etal.(1987)andArbiolandSa´ez(1988)describeda rodent-bearing site that theycalled Santpedor-2, placed350m eastfromtheclassicallocalityofSantpedor,andsituatedinthe same stratigraphic level (Fig. 1). These authors reported in Santpedor-2 thepresence of fourdifferent rodents:Theridomys Geobiosxxx(2019)xxx–xxx ARTICLE INFO Articlehistory: Received21January2019 Accepted21June2019 Availableonlinexxx Keywords: Rodentia EarlyOligocene Rupelian GrandeCoupure IberianPeninsula ABSTRACT

Thispaperprovidesarevisedandupdateddescriptionandidentificationoftherodentsfrom Santpedor-2(northeasternSpain).Theageofthislocalityisparticularlyrelevant,becauseitisoneofthefew EuropeansitesassignedtotheearlyOligocenethathasyieldedprimateremains.However,therodent assemblagefromthissitehasneverbeendescribedindetail,andthereforetheOligoceneageofthis localityremainedtentative.Despitethescarcityofthematerial,therodentsfromSantpedor-2are characteristicenougheithertobeidentifiedasalreadyknownspecies,oratleastcomparedwithclose species.Eightdifferentrodenttaxawererecognized.Amongthem,thetheridomyidEctropomysexiguus, whichisrecordedonlyinlateEocenelocalities,andthegliridGlamysdevoogdi,whichisknowninboth latestEoceneandearlyOligocenelocalities.TheothertaxaarefoundonlyinOligocenelocalities,and consistofthetheridomyidParatheridomysmargaritaeand membersoftwofamiliesthatarrivedin EuropeatthebeginningoftheOligocene:eomyids(Eomyssp.)andcricetids(Eucricetodonatavus,cf. Heterocricetodonsp.,cf.Pseudocricetodonsp.).Finally,anothertheridomyid(Issiodoromyinae)seemsto correspondtoanewspeciesandnewgenus,themainfeaturesofwhichcorrespondtoearlyOligocene evolutionarygrades.ThisrodentassemblagefromSantpedor-2isassignedtotheearlyOligocene,mainly onthebasisofthepresenceofimmigrantrodentsthatreachedEuropeinrelationwiththeGrande Coupure(cricetidsandeomyids).This datingisnotablysupportedbythepresenceofP.margaritae togetherwithGlamysdevoogdi,bothbeingfoundinHoogbutsel(earlyRupelian,MP21).Onlyonerodent (E.exiguus)representsanEocenerelict,asisthecaseofPseudoloris,theonlyprimaterecordedinthis locality.Therefore,thisstudysupportstheOligoceneageofSantpedor-2andthenthepersistenceof omomyidprimatesintheIberianPeninsula(atleastlocally)aftertheGrandeCoupure.

§

Correspondingeditor:GillesEscarguel. * Correspondingauthor.

aff.aquatilis,Gliravusaff.priscus,Eucricetodonatavus,and Pseudo-ltinomysgaillardi.Basedonthisrodentassemblage,theyassigned thelocalitytotheearlyOligocene,anagemaintainedinseveral laterworks(Cuencaetal.,1992;Aguilaretal.,1997;Ko¨hlerand Moya`-Sola`,1999;Barbera` etal.,2001;Hookeretal.,2007;Hooker, 2010;Minwer-Barakatetal.,2010,2012,2013,2015;Costaetal., 2011;Marigo´ etal.,2014;Sanjuanetal.,2014).Evenmore,Agustı´ etal.(1987)proposedabiozonebasedontherodentassemblage fromSantpedor-2.TheydefinedtheTheridomysaff.aquatiliszone, which would represent the earliest Oligocene mammal level (MP21),therebybeingcorrelatedwiththelocalityofHoogbutselin Belgium.However,neitherAgustı´ etal.(1987)norArbiolandSa´ez (1988)provideddescriptions,measurementsorillustrationsofthe rodentmaterialfromSantpedor-2,andtheirtaxonomicascriptions remainednonverifiableasaresult.

The relevance of this locality increased when Ko¨hler and Moya`-Sola` (1999) reported the presence of scarce primate

remains in Santpedor-2 and in the nearby and somewhat younger locality of Fonollosa-13. The fossil primates from Santpedor-2 consisted of two fragments of lower molars, whereasthosefromFonollosa-13documentedanuppertooth row,thedentalmorphologyofwhichallowedtoundoubtedly assign them to the genusPseudoloris. Interestingly, primates were so far supposed to have disappeared from Europe in relationtothedrasticcoolingrecordedattheEocene–Oligocene transitionandthesubsequentfaunalturnoverthatdramatically affected continentalvertebrate fauna, known as the ‘‘Grande Coupure’’.ThefindingofPseudoloris(P.godinoti)from Santpe-dor-2 and Fonollosa-13 represented the unique record of Oligoceneprimatesinthecontinent,apresencethatwasshortly afterconfirmedbythediscoveryofasinglelowermolarofthe genusMicrochoerusintheearlyOligocenelocalityofAguato´n, also in Spain (Pela´ez-Campomanes, 2000). However, the age proposed for Santpedor was based only on the faunal list

Fig.1.GeographicandgeologicalsettingoftheSantpedor-2fossillocality.A.SituationofthestudiedsectionintheIberianPeninsula.B.Detailedmapshowingthepositionof Santpedor-2(modifiedfromArbiolandSa´ez,1988).C.SimplifiedcolumnoftheSantpedorsectionindicatingthestratigraphicpositionofthesampledlevel(modifiedfrom

Costaetal.,2011).

M. Vianey-Liaudetal./Geobiosxxx(2019)xxx–xxx 2

providedbyAgustı´ etal.(1987)andArbiolandSa´ez(1988)and, despitethefactthattheinterestofthelocalityliespreciselyon its Oligocene age, a detailed study of its rodents was never developed.

Inthispaperwepresentathoroughstudyoftherodentsfrom Santpedor-2.Therodentassemblagefromthissiteincludeseight differentformsoftheridomyids,eomyids,cricetidsandglirids,and isthereforemorediverse thanpreviouslythought.Thedetailed studyof theserodentssupports theearly Oligoceneageof the locality.

2. Geologicalsetting

TheEbroBasinisthesouthernforelandbasinofthePyrenean Range,whichwasformedinresponsetoconvergenceandcollision betweentheIberianandEuropeanplatesfromtheLateCretaceous totheMiocene(Puigdefa`bregasetal.,1992;Busquetsetal.,2003). Thisbasinisdelimitedbythreemountainrangesformedduring theEocene-Oligocenecompression:thePyreneestotheNorth,the IberianRangetotheSouthwest,andtheCatalanCoastalRangesto the Southeast. The marine connection of the Ebro Basin was interruptedduringthePriabonian(themarine-continental transi-tionhasbeenpreciselydatedat36.0MabyCostaetal.,2010). FromthelateEocenetothelateMiocene,theEbroBasindisplayed a closed drainage system, and its depositional framework consistedofdistributivealluvialsystemsfedfromthetectonically activesurroundingranges,laterallychangingtoshallowlacustrine zones(Agustı´ etal.,1987;Barbera` etal.,2001;Sa´ezetal.,2007). Thebasininfill(upto2000mthick)consiststhereforeofalluvial, fluvialandlacustrinedeposits,inwhich severalvertebratesites have been identified (Agustı´ etal., 1987; Anado´n et al., 1987; Cuencaetal.,1992).Nevertheless,mammalsitescorrespondingto theearliestphaseofendorheicsedimentationarerelativelyscarce: theonlytwo localities assignedto thelateEocene (Priabonian, MP19/20) are Sant Cugat de Gavadons (Crusafont-Pairo´, 1967; Hooker et al., 2009; Minwer-Barakat et al., 2013, 2016) and RocafortdeQueralt(Anado´netal.,1987),whilethefirstmammal sitethat,accordingtothepublisheddata,postdatesthe‘‘Grande Coupure’’isSantpedor(Ko¨hlerandMoya`-Sola`,1999;Barbera` etal., 2001;Costaetal.,2011).

3. Materialandmethods

ThedescribedrodentspecimensfromSantpedor-2consistof 33isolatedteeth,housedinthecollectionsoftheInstitutCatala` de Paleontologia Miquel Crusafont (Sabadell, Barcelona, Spain). The nomenclatureused in thedescriptions (Fig. 2)follows Vianey-LiaudandMarivaux(2017)fortheridomorphs,Dienemann(1987) forcricetids, Vianey-Liaud(1994) for glirids,and Maridet etal. (2010)foreomyids.Measurements(Table1)weretakenusingan opticcalliper‘‘Nikonmeasuroscope10’’connectedtoamonitor ‘‘Nikon SC112’’; measured parameters include the mesiodistal maximumlengthandthemaximumwidthperpendiculartothe length,takeninkeepingthewearsurfacehorizontal.Micrographs wereobtainedusingtheEnvironmentalScanningElectron Micro-scope(ESEM)attheUniversitatdeBarcelona.

4. Systematicpaleontology ClassMammaliaLinnaeus,1758 OrderRodentiaBowdich,1821 FamilyTheridomyidaeAlston,1876 SubfamilyOltinomyinaeHartenberger,1971 GenusEctropomysBosmaetSchmidt-Kittler,1972 EctropomysexiguusBosmaetSchmidt-Kittler,1972

Fig.3(A–C)

Material:DP4:IPS105269;p4:IPS105286,m2:IPS105287. Description:

Upperteeth.TheuniqueuppertoothisaDP4(Fig.3(A)),the lingual sideof itscrownbeingrelativelyhighandnarrowwith respect to the buccal one; it is not anteriorly reduced. The postprotocristamakesthehighendoloph,whichdoesnotjointhe hypocone,andthenthesynclineIIcommunicateswiththeshallow andnarrowsinus.Thehighanterolophjoinstheparaconetothe protocone, closing the syncline I (anteroflexus) buccally. The continuous protoloph joins the apex of the protocone. The mesoflexus (syncline II+syncline I) is closed buccally by the mesostyleandtheextensionsofthepostparacristamesiallyandof thepremetacristadistally.Themesolophislowerthantheother lophsandbreaksatmid-length.Itsbuccalpartweaklyattachesits lingualpart, whichfuseswiththethickerlingual metaloph.The mesoloph separatesthe longbuccolinguallysyncline IIand the shortersynclineIII.Thebuccalmetalophistransversebutslightly mesially oriented, connected to the lingual metaloph, and separatedfromthelongposteroloph.Asaresult,thesynclineIV ismoredevelopedthanontheknownpopulationsofE.exiguusor E. monacensis.ThissynclineIVcouldbesubdividedintoamuch wornDP4,sinceaverylowbeadisvisibleatitsbottombothonthe distalflankofthebuccalmetalophandonthelingualflankofthe posteroloph.

Lowerteeth.Thep4fromSantpedor-2(Fig.3(B))ismoreworn thanthatofRocafortdeQueralt,thefrontpartofthelatterbeing incomplete.Itshowsabreakfromtheentoconidtothesinusid.On thisp4,theprotoconidandthelargermetaconidarelinkedbya completetransversemetalophulidI.Thelatterisnotchedmesially byashortantesinusid,limitedmesiallybyalowanterolophid.A high postmetacristid joins the mesostylid. The mesostylid is connectedtoamesolophidthatreachesthemesoconidswellingat the distal end of a strong postprotocristid. The synclinid II is divided by a short mesiodistal cristid, extending from the mesolophidtothemetaconid.ThesynclinidIIIisclosedlingually. The short distal ectolophid is separated by a notch from the mesoconidswelling. Theentolophidandposterolophidarehigh andcomplete,betweenthestrongentoconidandhypoconid.The morphologyofm2(Fig.3(C))isreminiscenttothatofthetooth fromSanCugatdeGavadons,inwhich thesmallanteroconidis framed by the anterolophid and anterocingulid, which delimit narrowanteroflexidandantesinusid.Thesinusidisslightlydeeper transversely, and the mesolophid is complete, reaching the mesostylid-postmetacristidjunction.

Remarks:TheteetharelargerthanthoseofEctropomysexiguus from the Isle of Wight, Rocafort de Queralt and San Cugat de Gavadons,but theyareclearly smallerthanOltinomysplatyceps (BosmaandSchmidt-Kittler,1972;Vianey-Liaudetal.,1994).The general features oftheseteetharecompatible withthespecies E. exiguus,whereasthefewdifferencesaremorederivedthanin thematerialfromtheabovementionedlocalities(largersize,much more markedlophodonty,highercrown,deeper sinusid).These characteristics argue for a form younger than the late Eocene populationsofE.exiguus.

SubfamilyTheridomyinaeAlston,1876

GenusParatheridomysVianey-LiaudetMarivaux,2017 Paratheridomysmargaritae(Vianey-Liaud,1972) Fig.3(D–F)

Material:M2:IPS18941;M3:IPS105257;m3:IPS105284. Description:

Upper molars.Themesialanddistalflanksof thecrownare relativelyflat,verticalandparallelonM2(Fig.3(D)).Thistoothis worn,andthewearreducestheposterosyncline(synclineIV)toan islet.Theparacone,mesostyleandmetaconeareslightlybulged.

Fig.2.DentalnomenclatureemployedforthedescriptionoftheteethofTheridomyidae(A,B),Cricetidae(C–F),Eomyidae(G)andGliridae(H).A,C,GandHrepresentm1;B andErepresentM1;Dcorrespondstoam2,andFtoaM2.AandBaremodifiedfromBonilla-Salomo´netal.(2016);GismodifiedfromDaxner-Ho¨ckandHo¨ck(2009);therest offigurescorrespondtooriginaldrawings.Abbreviationsforlowermolars:aaprcd:anteriorarmofprotoconid;abcld:anterobuccalcingulid;acd:anteroconid,anld: anterolophid;anlpd:anterolophulid;antsd:anterosinusid;buald:buccalanterolophid;eclpd:ectolophid;ecmslpd:ectomesolophid;ecstd:ectostylid;encd:entoconid; enlpd:entolophid;ensd:entosinusid;hyd:hypoconid;hyld:hypoconulid;lc:longitudinalcrest;linald:lingualanterolophid;mecd:metaconid;melpd:metalophid;melpdI: metalophidI;melpdII:metalophulidII;mescd:mesoconid;mestd:mesostylid;metstd:metastylid;mslpd:mesolophid;papcrd:posteriorarmofprotoconid;prcd: protoconid;prhycd:prehypocristid;prsd:protosinusid;psthyd:posteriorarmofhypoconid;pstlpd:posterolophid;pstmecd:postmetacristid;pstprcd:postprotocristid;sd: sinusid;sydI:synclinidI;.sydII:synclinidII;sydIII:synclinidIII;sydIV:synclinidIV.Abbreviationsforuppermolars:ac:anterocone;alp:anteroloph;alpl:anterolophule; asin:anterosinus;ast:anterostyle;bucalp:buccalanteroloph;bucg:buccalcingulum;endlp:endoloph;ensin:entosinus;enst:entostyle;hy:hypocone;linalp:lingual anteroloph;lincg:lingualcingulum;me:metacone;mesc:mesocone;mesin:mesosinus;mesl:mesoloph;mest:mesostyle;metcl:metaconule;metlp:metaloph;metlpl: metalophule;metst:metastyle;mu:mure;pa:paracone;pacl:paraconule;past:parastyle;prc:protocone;prhyc:prehypocrista;prl:protoloph;prlp1:protolophule1; prlp2:protolophule2;prmec:premetacrista;prprc:preprotocrista;prsin:protosinus;prst:protostyle;psc:posterocone;pspac:postparacrista;pstl:posteroloph;pstsin: posterosinus;syI:synclineI;.syII:synclineII;syIII:synclineIII;syIV:synclineIV.

Theprotoconeis notstronglypinched,and ratherappearsonly slightlystretchedobliquely(mesiobuccaltodistolingual),giving thesameorientationtothesinus.ThesynclineIandsynclineIIare equallylongandshorterthansynclineIII.Theenamelofthedistal flankoftheprotoconeisonlyalittlethinnerthanonthemesial flank;theenamelofthehypoconeandofitsarmsisparticularly thick,andasthickasthedistalborderofthecrown.

Lower molar. Only one m3 can be referred to this species (Fig.3(F)).ThereisnosynclinidI;themetalophidIismesialandthe synclinidII is notcompressed. The postmetacristidis highand joinsthe mesostylidat the lingual extremity of the transverse mesolophid.Thepostprotocristidisnotmarkedlyobliqueandit joinsthe(pseudo)mesoconidatthemesialextremityoftheshort andslightlyobliquebuccomesialtodistolingualshortectolophid. ThesynclinidIIisnearlyaslongasthesynclinidIV,butwider. SynclinidIIIandsynclinidIVareequalinwidth.

Remarks:Thecharactersoftheuppermolarsdescribedabove are found on theM2 of the type population from Hoogbutsel (Belgium, MP21; Vianey-Liaud,1972, 1989). These teeth differ fromthose of Blainvillimys langeiin their moreregular enamel thickness, their lower crown, less pinched protocone and less obliquesinus.Thefeatures ofthem3described abovearevery similartothoseofthem3ofP.margaritaelackingsynclinidIfrom Hoogbutsel(Vianey-Liaud,1972).Them3fromSantpedorislower crowned,withamoreregularenamelthicknessandlessoblique posprotocristidthanthem3ofB.langei.TheteethfromSantpedor differfromTheridomysaquatilisintheirsmallersize.

?IssiodoromyinaeTullberg,1899 ?Issiodoromyinaenov.gen.,nov.sp. Fig.3(G–Q)

Material:P4:IPS105276;M1-2:IPS105275,105277,105278; M3:IPS 105279,105280;dp4: IPS18940;p4:IPS105281;m2: IPS105282;m3:IPS105283,105285.

Description:

Upperteeth.ThecrownsoftheP4andM1-2,andprobablyof thewornM3,showaconstrictionoftheoutlineatthelevelsofthe terminationoftheanteriorarmoftheprotocone(atthejunction withtheanteroloph)andoftheposteriorarmofthehypocone(at itsjunctionwiththeposteroloph).Theseshrinkings,welldistinct on thelesswornteethonly,areneverobservedontheM1-2of Paratheridomysmargaritae,Theridomysaquatilis,andBlainvillimys langei,aswellasonM1-2ofanyspeciesofPseudoltinomys.Onthe P4andM1-2,theparaconeandmetaconearemarkedlybulgedand theprotolophandmetalopharethick.Themesolophisthinnerand lowerthantheselophs.OnP4(Fig.3(G)),theanterolophisshort buttheanterosyncline(synclineI)islongerthaninthelateEocene andOligocenespeciesofPseudoltinomys.Theposterolophisshort, and the posterosyncline is particularly reduced, even on the weakly wornM1-2 IPS 105275 (Fig.3(H)).The sinusis shorter buccolinguallythantheoppositesynclineII,orhassimilarlength inthemostwornteeth.ContrawhatisseenonPseudoltinomys,the sinus and the syncline II are not communicating: they are separated by a very short mure as seen on the weakly worn M1-2(Fig.3(H)).

Lower teeth. Thedp4 (Fig.3(M)) is slightlyshorter than p4 (Fig.3(N)),andthelatterissimilarinlengththenthatofthem2and m3. On dp4, the weak protoconid and the following post-protocristid are loweredby wear, and then themetaconid and entoconid remainhigher.There is a verylow mesiodistal ridge descendingfromthemetaconidtothemiddleofthemesolophid. The latter is complete, and joins the postmetacristid at the mesostylidlevel.ThelingualopeningofthesynclinidIIIisnarrow. The sinusid is buccolingually short. The long and curved posterolophidisthickenedatthehypoconulidlevel,linguallyto the obliqueposthypocristid. On theunworn p4, themetaconid connectstotheprotoconidbyahighmesialmetalophid.Thereis noantesinusid.Themesolophidisslightlylowerthanthe meta-and ento-lophids.Themesolophid isbroken beforejoiningthe mesostylidlevel by a short and narrownotch. Thesynclinid II displays a short and weak mesiodistal ridge. The ectolophid is short,showingaveryshallowandnarrowbreakinitsmiddle.The synclinidIIIislinguallyopen.Thesinusidisbuccolinguallyshorter thanthesynclinidIII.Theentolophidisashighastheotherlophids, and theposterolophid is long, withoutdistinct swelling at the hypoconulidlevel.

The m2(Fig.3(O))is worn and theenamelshows a similar thicknessonthemesial,buccalanddistalsurfacesofthecrown. Thereisanindicationofthepresenceofanantesinusidasanangle onthemesialfaceofthemetalophulidI.ThesynclinidIIisdivided byamesiodistalridge.Thesinusidisobliqueanddirectedtothe buccalendofthesynclinidIV.ThesynclinidIIisclosedlingually, thesynclinidIIIopen,andthesynclinidIVshallowlyandnarrowly open.Them3IPS105283(Fig.3(P))displaysthesamefeaturesas those of them2,and, as it is more worn,the synclinidIII and synclinidIVareweaklyclosedlingually.Averynarrowbreakis observedbetweentheobliquesinusidandthesynclinidIV.Asfor them2,anangleofenamelon themetalophulidIindicatesthe presenceofa possibleantesinusid.Itiswellmarkedonthem3 IPS105285(Fig.3(Q)).

Remarks: This material documents the best represented species of theridomyid from Santpedor-2. It was previously referredtoPseudoltinomysgaillardi(seeSection1).Nevertheless, these teeth show strong differences with this species, and moreover, with the genus Pseudoltinomys, like the absence of break of the longitudinal crest on upper teeth, the reduced antesinusidortherelativelylargerp4.Itwouldbenecessarytoget morematerial todefinea newspecies, asitappearstobe,and probablyalsoanewgenus.Theteethhavebeendirectlycompared withseveraltheridomyidspeciesfromthelateEoceneandearly

Table1

Measurements(inmm)oftherodentteethfromSantpedor-2.

Taxon Cataloguenumber Tooth Length Width Ectropomysexiguus IPS105269 DP4 1.66 1.86

IPS105287 m2 1.91 1.80 IPS105286 p4 1.75 1.44 Paratheridomysmargaritae IPS18941 M2 1.94 2.63 IPS105257 M3 1.96 2.23 IPS105284 m3 – 1.72 Issiodoromyinaenov.gen.,novsp. IPS105276 P4 2.23 2.69 IPS105275 M1-2 2.16 2.72 IPS105277 M1-2 2.03 2.75 IPS105278 M1-2 – – IPS105279 M3 2.20 2.69 IPS105280 M3 2.03 2.42 IPS18940 dp4 2.14 1.41 IPS105281 p4 2.35 1.97 IPS105282 m2 2.39 2.07 IPS105283 m3 2.30 1.93 IPS105285 m3 – – Eomyssp. IPS105265 p4 1.08 1.03 IPS105262 m1-2 – 1.09 Eucricetodonatavus IPS105268 M1 1.92 1.23

IPS105266 M2 1.32 – IPS105264 M3 – – cf.Heterocricetodonsp. IPS105273 M1 – – IPS105258 M2 1.50 1.48 IPS105260 M2 1.51 -IPS105267 M3 1.18 1.24 IPS105259 m1 1.72 1.23 IPS105261 m2 – – IPS105263 m3 1.58 1.33 cf.Pseudocricetodonsp. IPS105274 m2 – 1.19 Glamysdevoogdi IPS105272 p4 0.93 0.86 IPS105270 m2 1.01 1.11 IPS105271 m2 – –

Fig.3.ESEMpicturesoftheteethofTheridomyidaefromSantpedor-2.A–C.EctropomysexiguusBosmaetSchmidt-Kittler,1972.A:rightDP4(IPS105269);B:rightp4 (IPS105286);C:leftm2(IPS105287).D–F.Paratheridomysmargaritae(Vianey-Liaud,1972).D:rightM2(IPS18941);E:rightM3(IPS105257);F:rightm3(IPS105284).G– Q. ?Issiodoromyinaenov.gen.,nov.sp.G:leftP4(IPS105276);H:rightM1-2(IPS105275);I:rightM1-2(IPS105277);J:rightM1-2(IPS105278);K:leftM3(IPS105279);L: leftM3(IPS105280);M:rightdp4(IPS18940);N:rightp4(IPS105281);O:rightm2(IPS105282);P:leftm3(IPS105283);Q:rightm3(IPS105285).Scalebar:3mm.

OligocenelocalitiesofSpainand SouthernFrance:SanCugatde Gavadons,CalafandLosBarros(EbroBasin,Spain);Civrac, Saint-Capraise-d’Eymet and Soumailles (Aquitaine Basin); Escamps, Sainte-Ne´boule, Aubrelong 1 and Ravet (Quercy Phosphorites); Ronzon(Auvergne).ThesizeislargerthanthatofPseudoltinomys aff. cuvieri from San Cugat de Gavadons and P. gaillardi from Soumailles,Ronzon,orfromtheQuercylocalities(Vianey-Liaud, 1976). The deciduous lower molar is simpler than that of P. gaillardi; it is less hypsodont and simpler than that of P. amblesi(GarzonHeydtandLo´pezMartı´nez,1978).Thep4has its mesialwidth less reduced withrespect tothe distalwidth compared with p4 of P. cuvieri, P. gaillardi, Paratheridomys margaritaeand Blainvillimys langei. The crownis lowerthan in P. gaillardi.ComparedtothespeciesofTheridomys,itisclosetothe sizeofthesmallestteethofT.aquatilis,andsmallerthanT.golpei andT.calafensis(Anado´netal.,1987).Inaddition,thedp4andp4 aresimplerandshorterthaninTheridomys.

FamilyEomyidaeDepe´retetDouxami,1902 GenusEomysSchlosser,1884

Eomyssp. Fig.4(A,B)

Material:p4:IPS105265;fragmentofm1-2:IPS105262. Descriptionandremarks:Thesizeoftheseteethislargerthan that of Eomysantiquus, closetothat of E.molassicus(Engesser, 1987),andslightlysmallerthanthatofE.zitteli(Comteand Vianey-Liaud, 1989). However, somefeatures of thesetwo teethdiffer fromthelattertwospecies.Thep4ofEomyssp.(Fig.4(A))isless elongated,anditsmetaconidandprotoconidaremorerobustthan onE.zitteli.ContrarytoE.antiquus,E.molassicusandE.zitteli,inthe teeth from Santpedor the anteroconid/anterolophid is absent: thereisonlyahighmesialconnectionbetweentheprotoconidand the metaconid,which is themetalophulid I. These cuspids are closertoeachotherthanonE.molassicus,onwhichthe‘trigonid’ appears less short buccolingually than on the species from

Fig.4.ESEMpicturesoftheteethofEomyidae,CricetidaeandGliridaefromSantpedor-2.A,B.Eomyssp.A:rightp4(IPS105265);B:rightm1-2(IPS105262).C–E.Eucricetodon atavusMisonne,1957.C:rightM1(IPS105268);D:rightM2(IPS105266);E:rightM3(IPS105264).F–L.cf.Heterocricetodonsp.F:leftM1(IPS105273);G:leftM2 (IPS105258);H:rightM2(IPS105260);I:leftM3(IPS105267);J:rightm1(IPS105259);K:leftm2(IPS105261);L:rightm3(IPS105263).M.cf.Pseudocricetodonsp.,leftm2 (IPS105274).N–P.Glamysdevoogdi(BosmaetdeBruijn,1979).N:leftm2(IPS105270);O:leftm2(IPS105271);P:rightp4(IPS105272).Scalebar:1mm.

Santpedor.Onp4andm2,themesolophidismoderatelylongas observedonE.zitteliorontheEomyssp.fromMo¨hren20(Maridet etal.,2010).ItislongerthanonthetypeofE.zitteli,butshorterand weakerthanonE.molassicus.Onthelatter(typefromOensingen), itmakesanangle,beingfirstlydirectedposteriorlybeforebeing transverse,whereasitistransversesincethemesoconidonEomys sp.fromSantpedor.Onthefragmentofm2IPS105262(Fig.4(B)), theattachment oftheentolophid ontheposterolophid ismore buccalthanonE.molassicus,andthustheposterosynclinidandthe posterolophidarelongermesiodistally.Therefore,thespeciesfrom Santpedordisplaysoriginalfeatures.

FamilyCricetidaeRochebrune,1883

Remarks: When studying the rare cricetid teeth from Santpedor-2, we were confronted with the absence or incom-pletenessofdiagnosisforseveralOligocenespeciesorgenera.The history of these European Oligocene cricetids, resulting from severalphasesofimmigrationandlocaldevelopmentsinWestern Europe,iscomplex.Anexhaustivereviewwithdirectcomparison ofpopulationsfromnumerouslocalitiesinthisgeographicalarea would be essential to achieve consistent results. It would be necessarytotakeintoaccountalltheirfeatures,dental (morphol-ogyandultrastructure)andcranial.

SubfamilyEucricetodontinaeMeinetFreudenthal,1971 GenusEucricetodonThaler,1966

EucricetodonatavusMisonne,1957 Fig.4(C–E)

Material:M1:IPS105268;M2:IPS105266;M3:IPS105264. Description:

M1. Theenameloftheoutskirtsofthecrowniswrinkled.The buccalsideisconvex.Thereisonlyasingleanterocone,fromwhicha weakanterocone spurslopes andjoins the long protocone spur. Lingually,theanterolophslopestotheleveloftheprotostyle,from whichitisseparatedbyanotch.Buccally,thecingulumislongerand strongerthantheanteroloph.Theprotolophulejoinstheshortdistal armoftheprotocone,andthemetalophulereachestheshortmesial armofthehypocone.Theentolophislongandanglesatthelevelof themesocone.Themesolophisshortandthemesostyleisreducedto shortcingularelements.Lingually,theentostyleisalsoreduced.

M2. The buccal anteroloph is well developed, reaching the buccalcornerofthetooth.Thelingualanterolophisthinner,lower and long. The protoloph is oblique forwardly and joins the anterolophule.Theposteriorarmoftheprotoconeispresent.The mesolophisshort.Themetalophuleisdirectedanteriorly.Athin entostyleunderlinesthebuccalborderofthemesosinus.Itdiffers fromtheM2ofcf.HeterocricetodonfromSantpedor-2initssmaller size,lophsandmesosinuslesstransverseandinthedifferentwear pattern(horizontalonlyon thelingual halfofEucricetodon, the paraconeandmetaconebeingbulgedandprotruding,whereasthe buccalcuspsofHeterocricetodonarealsoflattened).

M3.Thistinytoothistoomucherodedtodepictitsfeatures. Remarks:ThesmallspeciesofEucricetodon,includingatavus, havebeenreferredtothegenusAtavocricetodonbyFreudenthal (1996),butthisauthordidnotgiveanydifferentialdiagnosiswith respecttothegenusEucricetodon.Adifferentialdiagnosisisalso missing in the redescription of the species atavus made by Freudenthal (1988). The validity of Atavocricetodon was then discussedandrejectedbyotherauthors(e.g.,deBruijnetal.,2003, GomesRodriguesetal.,2013),who‘‘foundnoobviousreason,even pragmatical, to keep Atavocricetodon as a genus or even a subgenus’’. This is the reason why we decide here to use EucricetodonratherthanAtavocricetodon.Asthiscricetidispoorly represented in the locality, we cannot contribute more to this discussion. Its size and morphology fit those of the genus Eucricetodon:bunodontteeth,largerthanthoseof Pseudocriceto-donand withsimpler lophs and ridges; anterocone of theM1

generally simple, sometimes tending to be slightly duplicated, convexbuccalcontourof theM1and posteriorlobe of theM3 markedlyreduced(Vianey-Liaud,1972).Thesizeandmorphology oftheM1andM2matchthoseofthelowerOligocenepopulations ofE.atavus(e.g.,Hoogbutsel,Aubrelong1,Ravet,Montalba´n,Me`ge, Pech Crabit, Itardies; Vianey-Liaud, 1972, 1974; Freudenthal, 1988), better than to ‘Atavocricetodon’ aff. nanus from Valbro (Peigne´ etal.,2014:fig.16).Themoredistinctivedifferenceinsize is between thelowerm1 ofthesetwo species, but thereis no availablem1inSantpedor-2.Ifconsideringthetypepopulationof nanusfromValdecollares(Pela´ez-Campomanes,1995),thesizeof theM1andM2fromSantpedor-2isclearlylarger. Eucricetodon atavus is recorded from MP21 to MP23 (Vianey-Liaud, 1972; Vianey-LiaudandSchmid,2009).

SubfamilyPseudocricetodontinaeEngesser,1987 GenusHeterocricetodonSchaub,1925

cf.Heterocricetodonsp. Fig.4(F–L)

Material: M1 (distal part): IPS 105273; M2: IPS 105258, 105260;M3:IPS105267;m1:IPS105259;m2:IPS105261;m3: IPS105263.

Description: We refer the teeth from Santpedor to a small primitivespecies ofHeterocricetodonbetterthanto Pseudocrice-todon,onthebasisoftheirratherlophodontdentalpatternandthe presenceofalongmesolophononeoftheM2(Fig.4(G))andonthe M3 (Fig. 4(I)). Even if this latter feature is variably found in Pseudocricetodon incertus, it is observed in the material from SantpedortogetherwithotherfeaturesfoundinHeterocricetodon: thelonglingualanterolophofM2,thehighlophsontheunworn M3;theflatwearsurfaceoftheM2fromthelingualedgeofthe crown to the 2/3 of the occlusal surface, with moderately prominent buccal upper main cusps; the long postmetacristid on m1(Fig.4(J)), theflatwearsurfaceofthem3,withitslong posterior arm of the protoconid, and the reduced width of its posteriorlobe(Fig.4(L)).Theanteroconidandmetaconidofm1are stronglyworn,andthereforethelophidsaredifficulttodistinguish, excepttheoccurrenceofalongpostmetacristidandashortthick posteriorarmoftheprotoconid,weaklyseparatedfromthelingual metalophulid;therearetwoshortmesolophids,themoremesial beingthelongest.Thereisnoposteriorarmofthehypoconid.The fragmentofm2showsalongpostmetacristidendingina small swellingasametastylid;itsposteriorarmoftheprotoconidislong andobliquetowardsthebase,andendswithathickeningagainst themiddleofthebuccalflankofthemetaconid;themesolophid hassamelengththantheposteriorarmoftheprotoconid.Flatand wornonitsbuccaltwo-third,itslingualcuspidsarenotworn.The enamelofthesurroundingsofthecrowniswrinkled.

Remarks:ThegenusHeterocricetodonwassofarunknownin thelowerOligocene.Untilnow,theearliestspeciesreferredtothis genuswereH.landroveri(Daamsetal.,1989)fromPareja(MP26, earlyChattian;Aguilar etal.,1997)andH.hausifromBumbach (MP24-25?,earliestChattian;Engesser,1987).

AsnoticedbyEngesser(1987:p.990)forthespecieshausi,itis difficulttoreferisolatedteethtoHeterocricetodon,andespeciallyto differentiate them from the genus Pseudocricetodon until the discoveryofmoreabundantmaterial.Itisevenmuchmoredifficult tocarryoutsuchadistinctioninanolderRupelianlocality,where veryfewteetharedocumented.Daamsetal.(1989:pp.47–48) proposetoincludethespeciesincertuswithinthe genus Hetero-cricetodonevenifcranialfeatures‘‘provethevalueofthisgenus’’. The teeth from Santpedor are largerthan those of ‘Pseudo-Pseudocricetodon’ incertus from some lower Chattian localities (Vianey-Liaudetal.,2014:table5,p.594)butsmallerthanH.hausi fromBumbach,andsimilarinsizetoH.landroveri(m1smallerthan thesmallestm1fromPareja,m3likethelargest).Thespeciesfrom

Santpedor-2 differs from H. landroveri in the more reduced posterior lobe of m3 and themesoconid more distinct on m1. ItsmorphologyisclosertoH.hausi,butthesinusseemslessdeep buccolingually. As the sample is scarce, we cannot define the morphological and size variability, and prefer to keep this Heterocricetodon sp. in open nomenclature. This species thus representstheearliestrecordofthegenusdocumentedsofar.

GenusPseudocricetodonThaler,1969 cf.Pseudocricetodonsp.

Fig.4(M)

Material:Anincompletem2:IPS105274.

Descriptionandremarks:Thisfragmentoftoothdiffersfrom thatascribedtoHeterocricetodoninitsslightlysmallersize,the enamelsmooth,andthemorebulbousandroundedmaincuspids, aswellasinthedifferentarrangementofthetalonidbasinarea. Thepostmetacristidisshorterandthemetastylidlessdistinct,by contrast there is a mesostylid blocking the opening of the entosinusid.Thereis no mesolophiddistinct fromtheposterior armoftheprotoconid.Thelatteriswidelyobliqueandisconnected tothemesialextremityoftheectolophid:atthejunctionitcurves buccolingually.If this interpretation is right,themesolophid is absent,becausethereis nodistinctmesoconid,and no ectome-solophid,butonlyafaintverticalswellingonthebuccalflankofthe shortectolophid.

FamilyGliridaeThomas,1897

SubfamilyGlamyiinaeVianey-Liaud,1994 GenusGlamysVianey-Liaud,1989

Glamysdevoogdi(BosmaetdeBruijn,1979) Fig.4(N–P)

Material: p4:IPS 105272;m2: IPS 105270;m2 incomplete: IPS105271.

Description:Thewearsurfaceofp4(Fig.4(P))isnearlyflatand thecrownrelativelylow.Themetaconidandprotoconidarelinked bytwo shortand thicklophids:ananterolophidand a metalo-phulidII.Theanteriorarmofthehypoconidisfaintlydeveloped; therefore,thesinusidisconfluentwiththeposterosynclinid.The posterolophid is long and strong, ending against the bulged entoconid.On m2,theanterolophidis long,separates fromthe protoconidandjoinsthemetaconid;thepostmetacristidispresent and slopes gently to the mesoflexid lingual opening; the metalophid is transverse and thick; the mesoconid is present, andthemesolophidislong,connectedtotheentoconidbyalow entolophid.Oneextra-ridge(Fig.4(N))andgranules(Fig.4(O))are presentinthemesoflexidandposteroflexid.

Remarks:Thesizeofthep4andofthecompletem2fitsthesize range ofGlamys devoogdi fromHamstead Bedsand Hoogbutsel (Bosma and de Bruijn, 1979; Vianey-Liaud, 1994) and the morphologyisalsoclosetothatofthisspecies.Onthep4figured inBosmaanddeBruijn(1979:pl.2,fig.5),themesialconnection (anterolophid)islowanddoesnotreachthetopofthemetaconid andprotoconid,whereasthisconnectionishigherandcompleteon thep4fromSantpedor.Thepathsoftheotherextra-ridges and granules,intothesinusidandthetalonid,occupysimilarpositions buttheirconnectionsvary.Thespeciesisrecordedfromthelate Priabonian(MP19/20)totheearlyRupelian(MP21).

5. Discussion

TheEocene-Oligocenetransitionwascharacterizedbyadrastic decrease in global temperatures, which involved a significant changeinthefaunasofbothmarineandterrestrialrealms,known as the ‘‘Grande Coupure’’ (e.g., Stehlin, 1910; Prothero, 1994; Zachosetal.,2001;Hookeretal.,2004;Hooker,2010;Hrenetal., 2013).IntheEuropeancontinentthisfaunalturnoverwasmarked,

amongotherchanges,bythearrivalofsomeimmigrantgroups, suchascricetidrodents(Vianey-Liaud,1979;Hookeretal.,2004; MaridetandNi,2013;FreudenthalandMartı´n-Sua´rez,2016),and bythenearlycompletedisappearanceofprimates.Untiltheendof theXXth century,it wasgenerally acceptedthat microchoerine primatesbecameextinctattheEocene-Oligocenetransition,but theidentificationofscarceremainsofPseudolorisinFonollosaand Santpedor (Ko¨hlerandMoya`-Sola`, 1999)and ofMicrochoerusin Aguato´n(Pela´ez-Campomanes,2000)testifiedtothepersistenceof thisgroupintheIberianPeninsuladuringtheearliestOligocene. Forthisreason,thedetailedstudyofthefossilsitesdocumenting this time window in Spainis particularlyrelevant, in order to ensureprecisedatingofsuchlocalities,andtoanalyzethemammal assemblages thatcoexisted withtheselast surviving Paleogene primatesinEurope.

The reference faunas of the latest Priabonian (MP20, St-Capraise-d’Eymet)andearliestRupelian(MP21,Soumailles) MP-levelsarepoor,andhavebeenchosenmainlybecausetheyhave providedbothsmallandlargemammals.However,the evolution-arystagesofthedocumentedlineagesarenotverydiscriminating, eitherfor theMP20or fortheMP21 (Aguilar et al.,1997). The characterization of the MP20, which is mainly based on the terminal stage of Palaeotherium curtum (sub-species frohnstet-tense), is not evident (e.g., Hooker, 2010). For the rodents, ‘Theridomys’ bonduellicould bea good marker, but this species is onlyknownfromtheParisand Hampshirebasins.Thishasa definiteimpactontheprecisedefinitionandcorrelationsoffaunas aroundthe‘‘GrandeCoupure’’.

The‘‘GrandeCoupure’’wouldmarkthebaseoftheRupelian, currentlydatedat 33.9Ma(Vandenbergheetal.,2012).Several correlation attempts withrespectto mammalianlineages have beenmade,usingmagnetostratigraphyandgeochemistry(Solent Group,WhitecliffBay,IsleofWight, England;Galeet al.,2006; Hookeretal.,2009)oronlymagnetostratigraphyinthecaseofthe Santpedor section (Barbera` et al., 2001; Costa et al., 2011). Accordingtothelatterauthors,themagnetostratigraphiclocation oftheMP21level,possiblyrepresentedatSantpedorattheupper partofthe13rchron,woulddatethisMParound33.4Ma,butthis datum needs to be confirmed. This locality contains both immigrant rodents, such as the Cricetidae, as well as native species of Gliridae and Theridomyidae. The scarcity of rodent materialintheMP20andMP21referencelevels,combinedwith thegeographicaldifferentiationobservedbetweentheridomyoids at the beginning of the Oligocene, does not facilitate precise correlations. Nonetheless, it is easy to identify the arrival of immigranttaxaandalsotodistinguishtheevolutionarystagesof certainlineageswhentheyhaverepresentedbysufficientteeth.

TherearecurrentlyfewRupelianandearliestChattianMP,four innumber(MP21toMP24),overatotalstagedurationof6myr. However,basedon theevolutionarydegreesofseveral Therido-myoidealineages,atleastoneintermediatelevelMP23/24canbe insertedeven ifit hasnot yetbeenformallydefined (Fig.5).It wouldcorrespondtotheevolutionarylevelofthefaunafrom St-Martin-de-Castillon (Vaucluse), in particular to the species Protechimystruci(Hugueney,1994;Vianey-Liaud,1998; Vianey-Liaud and Marivaux, 2017). The standard locality of MP24 (Heimersheim)islocatedatthebaseoftheCyrenenMergel,and correlatedwiththeKasselerMeeressand,whichprovided nanno-planktonfromtheNP24area(Bahlo,1975:p.12).Thus,theageof MP24 is close to the Rupelian-Chattian boundary (28.3 Ma; Vandenbergheetal.,2012)withinthelowermostChattian.

The studyof therodentsfrom Santpedor-2reveals a higher diversitythanpreviouslythought.Evenifthenumberofrodent teethissmall,theyarecharacteristicenougheithertobeidentified asalreadyknownspecies,oratleastcomparedtomorphologically closespecies.Amongthem,onespeciesisknownonlyfromlate

Pleasecitethisarticleinpressas:Vianey-Liaud,M.,etal.,TherodentsfromSantpedor-2(EbroBasin,NESpain)confirmtheOligocene ageofthelatestprimatesfromthePaleogeneofEurope.Geobios(2019),https://doi.org/10.1016/j.geobios.2019.06.002

Eocenelocalities(MP18toMP20):theTheridomyidaeEctropomys exiguus(Vianey-Liaudetal.,1994).ThegliridGlamysdevoogdiis knownfromlatePriaboniantoearlyRupelianlocalities(MP19to MP21;Vianey-Liaud,1994:table2).Theotheridentifiedrodents belongexclusivelytoOligocenetaxa(Fig.5).OneTheridomyidae (Paratheridomysmargaritae)isknownfromthestandardlocalityof MP21 (Hoogbutsel,Belgium; Vianey-Liaud,1972), which is the typelocalityofthespecies,andfromLaPlante2(Quercy,France, MP22;Vianey-Liaud,1989). TheotherTheridomyidae (?Issiodo-romyinae)isprobablyanewspecies,andperhapsanewgenus;its features(e.g.,sizeofpremolars)betterindicateOligocenerather thanlateEoceneevolutionarygrades.Twoimmigrantfamiliesof the ‘‘Grande Coupure’’ are recognized: the Eomyidae and the Cricetidae,amongwhichonespecies(Eucricetodonatavus)andone genus (Pseudocricetodon) are recorded since MP21. The other generaarerepresentedbyoriginalspecies,aHeterocricetodonmore

primitive,smaller, and plausibly olderthan theearliest species known until now, and an Eomys largerthan the MP21 species E. antiquus, but with features different from all known Eomys species.

ThisrodentassemblageismostprobablyearlyOligoceneinage, withacertaindiversityofwell-characterizedimmigrantfamilies, andOligoceneevolutionarygradesofautochthonouslineages(i.e., P. margaritae and G. devoogdi).One rodent species (Ectropomys exiguus)representsanEocenerelict,asisthecaseofPseudoloris godinoti,theonlyprimate recordedinthis locality.Asthereare only33teethforeightrodentspeciesfoundsofar,someofthem remaininopennomenclature,andthereforewecannotpreciseif the assemblage is close to MP21 or to MP22. Further field campaigns in this fossil site, leading to the recovery of more abundantmaterial,willprobablyallowformoreprecisetaxonomic determinationsandamoreaccuratedatingofthelocality.

Fig.5.KnownstratigraphicalextensionoftherodentsidentifiedinSantpedor-2,afterVianey-Liaud(1994),Freudenthal(1996),Hookeretal.(2004),Vianey-LiaudandSchmid (2009),Peigne´ etal.(2014),andVianey-LiaudandMarivaux(2017).Dottedlinesindicatetheextendedrangesofsometaxaaftertheidentificationoftherodentsfrom Santpedor-2(MP21orMP22).TheMPreferencelevelsarecorrelatedtotheGTSscale(Vandenbergheetal.,2012),withpositionoftheselevelsdiscussedinVianey-Liaudand Marivaux(2017).TheTheridomorphafromGousnat(MP18a)representevolutionarygradesmoreprimitivethanthosefromLaDe´bruge(MP18b)(Vianey-Liaudand Marivaux,2017).

6. Conclusions

Therodentfossilmaterial fromSantpedor-2(EbroBasin,NE Spain)wasdescribedindetailforthefirsttime.Theassemblage includes eight different rodent taxa: Ectropomys exiguus, Para-theridomysmargaritae,?Issiodoromyinaenov.gen.,nov.sp.,Eomys sp.,Eucricetodonatavus,cf.Heterocricetodonsp.,cf. Pseudocriceto-donsp., and Glamys devoogdi;it is notably differentfrom that reportedinthepreliminaryfaunallistsprovidedbyAgustı´ etal. (1987) and Arbiol and Sa´ez (1988). Although the locality has yieldedone species that can beconsidered as an Eocene relict (Ectropomys exiguus), the presence of two rodent families that arrivedinEuropeaspartofthe‘‘GrandeCoupure’’event(Eomyidae and Cricetidae),aswell astheevolutionarygrade of speciesof GliridaeandTheridomyidae,mightindicateanearlyOligoceneage for this assemblage. Nevertheless, the scarcity of the material preventsa precisetaxonomicdeterminationofsometaxaanda conclusiveassignmenttooneoftheearliest Oligocenestandard levels(MP21orMP22).Inanycase,thisstudyconfirmstheearly OligoceneageofSantpedor-2,oneofthefewEuropeanlocalities that documentsthepersistenceof omomyidprimatesafter the ‘‘GrandeCoupure’’.

Acknowledgements

We are truly grateful to the editor G. Escarguel and three anonymous reviewers for their constructive comments that greatlyimprovedthemanuscript.Thisworkhasbeensupported by the CERCA Programme/Generalitat de Catalunya, the project CGL2017-82654-P, MINECO/FEDER, UE (Ministerio de Economı´a, Industria y Competitividad, Spanish Government and European Union),andtheresearchgroups2017SGR86GRC(Generalitatde Catalunya)andRNM190(JuntadeAndalucı´a).R.M.-B.isfundedby thePlanPropio deInvestigacio´n,VicerrectoradodeInvestigacio´ny Transferencia,UniversidaddeGranada.Thisworkhasmadeuseof the collections of the Institut des Sciences de l’E´volution de Montpellier(ISE-M,UMR5554).Publ.ISEMno_2019-125.

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