Çeştepe, a new early Pliocene vertebrate locality in Central Anatolia and its stratigraphic context

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Originalarticle

C¸ es¸tepe, a new early Pliocene vertebrate locality in Central Anatolia

and its stratigraphic context

C¸es¸tepe, une nouvelle localité à vertébrés du Pliocène inférieur d’Anatolie centrale

et contexte stratigraphique

SevketSena,∗,MassimoDelfinob,c,NizamettinKazancid

aUMR7207CNRS,Muséum,UPMC,CentredeRecherchessurlaPaléobiodiversitéetPaléoenvironnements,8,rueBuffon,75005Paris,France
bDipar timen todiS cienzede llaTerr a,Unive rsi tàdiTorino ,Via T .ValpergaCaluso 35 ,10125Torino,Italy
cInstitutCatalà d ePaleon tolog iaMig uelCrusafo nt, Univer sitat A utònoma deBarc elo na,Edi ficiICTA -ICP,CarrerdelesColumness/n,CampusdelaUAB,

CerdanyoladelVallès,Barcelona08193,Spain
dAnkaraUn ive rsity,G eologicalE ngineer ingDepartment,Tandog˘an,Ankara06100,Turkey

a r t i c l e i n f o a b s t r a c t

Articlehistory: ThenewlocalityofC¸es¸tepeintheKazanBasin,45kmNNWofAnkara,yieldedtheremainsofthree
Receiv ed13November2016 spec ieso freptiles (cf .Lac erta sp. ,Ang uinae indet.a nd Var anussp .) andanoc hotonid lago morph,O ch otona
AAvccaeilpatbelde 2on5 lJianneuxaxrxy 20 17 mediter ran ensis Su ata Alpasla n, 2 009. The l atter s peci es sugge sts a n ea rly Pliocene a ge (MN14). The fossil
layerisincludedintheSinapFormationthatdeliveredabundantremainsofMio-Pliocenemammals

inthesouthernpartofthebasin,butinthenorthernpartofthebasinC¸es¸tepeiscurrentlytheunique
Keywords: fos sils ite.Thea geas sig ned bythe fau na led torevise thes tra tigra phyo f se dime nt aryforma tion sinthe
Lagomorpha
C¸es¸tepearea,previouslyconfusingbecauseofthevariouslithostratigraphicunitnamesandagesgiven
Ochotona
inpreviouswork.ThetaxadescribedinthisnotearestillpoorlyknowninTurkey.Varanushasbeen
Squamata
previouslyreportedfromC¸alta,alocality16.3kmSWtoC¸es¸tepeinthesamebasin,andO.mediterranensis
Varanus
EarlyPliocene fromIg˘deliintheSivasBasin.Althoughrepresentedbyafewtaxa,theC¸es¸tepefaunuleshowsEurasian
Kazan Basin affini ties.
Turkey ©2017ElsevierMassonSAS.Allrightsreserved.

r é s u m é

Motsclés: LanouvellelocalitédeC¸es¸tepedanslebassindeKazan,à45kmauNNWd’Ankara,alivrélesrestesde

Lagomorpha troisespècesdereptiles(cf.Lacertasp.,Anguinaeindet.andVaranussp.)etd’unlagomorpheochotonidé,
Ochotona Ocho tonamed ite rranensi sSu ataAlpa slan ,2009.Ce ttefau nee tenpart iculi er lader nièreespèce permettent
Reptilia decorréle rcettelocalitéa uPlio cèneinfér ieur( MN14 ).Lac ou ch efossilifère e stincluse dansla Formation
Varanus de Sinapqu ialiv réd’abo nda ntsreste sdemam mifères mi o-pliocè nesdansl apa rtiesud dece b assin,mais
Pliocèneinférieur
danslapartienorddubassin,C¸es¸tepeestpourlemomentl’uniquesitefossilifère.L’âgeattribuéparcette
BassindeKazan
fauneapermisderevoirlastratigraphiedesformationssédimentairesdanscesecteurdubassindeKazan,
Turquie
auparavantconfuseenraisondesnomsd’unitéslithostratigraphiquesetdesâgesdifférentsdansdes

travauxantérieurs.LestaxonsdécritsdanslaprésentenotesontencoremalconnusenTurquie.Legenre

Varanusn’aétédécritauparavantquedugisementdeC¸altaauSEdumêmebassinetO.mediterranensisdu

gisementd’Ig˘delidanslebassindeSivas.Bienquepauvre,cettefaunulemontredesaffinitéseurasiatiques.

©2017ElsevierMassonSAS.Tousdroitsre´serve´s.

1. Introduction

PlioceneterrestrialdepositsoccupyquitelargeareasinTurkey.
∗
Correspondingauthor.
However,Pliocenevertebratelocalitiesareratherrareandmuch
E-mailaddresses:[email protected](S.Sen),massimo.delfi[email protected]
(M.Delfino ),kazanci@ eng.ankara.edu.tr(N .K azanc i). lessdocumented.IntheirzonalsubdivisionofterrestrialNeogene

http://dx.doi.org/10.1016/j.annpal.2017.01.004
0753-3969/©2017ElsevierMassonSAS.Allrightsreserved.

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

context.AnnalesdePaléontologie(2017),http://dx.doi.org/10.1016/j.annpal.2017.01.004
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in Turkey, Ünay et al. (2003) suggested four mammalian zones TuzgölüTertiarybasins)aredeveloped.TheAnkararegionformsa

(M to P) for Pliocene based on rodent assemblages. However, kindofnodewherethesetectonicunitsandsedimentarybasinsget

thecurrentstateoftheknowledgeaboutthePliocenemammals closetoeachotherand/orimbricated(Fig.1).Toclarifysuchacom-

doesnotallowtheeffectiveuseofthissubdivision,neitheraccu- plexgeologicalcontextneedsrobuststratigraphic,biostratigraphic

rate correlations with the MN zonation. The main issue is that andanykindofchronologicaldata.Therefore,theC¸es¸tepefossils

the Pliocene mammal faunas of Turkey are still poorly docu- areofinterestnotonlyintermsofsystematicsandpaleoecologyof

mented and their composition is somewhat different from that speciestheyrepresent,butalsofortheircontributiontounderstand

of the surrounding regions. For Pliocene of Turkey, the best- thegeologicalevolutionoftheregionthatisincreasinglycomplex

documented fauna is probably C¸alta, at some 50km NW to throughouttheNeogene.

Ankara,inthesamebasinasC¸es¸tepe(Sen,1998a,1998b).Other The Gondwana-Eurasia collision, which gave rise to the

Pliocene localities, dispersed in sedimentary basins across all Alpine-Himalayan mountain chain, created the framework of

Turkey (Sarac¸, 2003), are mainly known with their rodent asso- the current geography of Anatolia. The Anatolian plate became

ciations,inparticularthosestudiedbyÜnayanddeBruijn(1998) uplifted during the Neogene, and several large and small basins

and Van den Hoek Ostende et al. (2015a, 2015b). The discovery appearedtoreceiveterrestrialsediments(Fig.1B).Inrelationwith

of a new locality at C¸es¸tepe (Kazan Basin) provided opportunity thecollision,extensivevolcanicprovincesdevelopedindifferent

to review stratigraphic correlations in this basin and to doc- regions of Anatolia since the early Miocene (e.g. East Anatolian

ument Pliocene reptiles and ochotonid lagomorphs in Turkey. VolcanicProvince–EAVP,CappadocianVolcanicProvince–CVP,

As we will see here below, these groups may provide strong KızılcahamamVolcanicProvince–KVP)and,asexamplifiedinthis

arguments for biochronology and environment if they are well- study,volcanismimpactedterrestrialinfilloftheseNeogenebasins

documented. (Figs.1and2).Localdifferencesinvolcanic,volcano-sedimentary

and sedimentary input in these basins produced stratigraphic

successionsthatareheterogeneousanddifficulttocorrelateeven

2. Geologicalcontext insidethesamebasin.ThisisparticularlytrueforNeogenedeposits

ofCentralAnatolianbasins(Tatlı,1975;Öngür,1976;Göktenetal.,

2.1. GeologicalcontextofCentralAnatoliaandKazanBasin 1988; Türkecan et al., 1991; Duru and Aksay, 2002). The North

AnatolianFaultZone(NAFZ),whichisthesecondlongeststrike-slip

The C¸es¸tepe village and vertebrate fossil locality are situated fractureoftheworldaftertheSanAndreasFault,issituatedafew

about45kmNNWtothecapitalAnkara,and14kmnorthofKazan kilometersnorthofthestudyarea,anditseparatestheEurasian

townintheKazanBasin,CentralAnatolia(Fig.1).Thisbasinisat and Anatolian plates (Fig. 1). The occurrence of NAFZ indicates

thesouthernmarginoftheKızılcahamamVolcanicProvince(KVP thebeginningoftheneotectonicperiodinAnatolia,anditisoften

inFig.1A)whichcoverslargeareasNWtoAnkaraandsouthtothe consideredthesourceofincreasingseismicactivity(Sengöretal.,

NorthAnatolianFaultZone(NAFZ).Inpreviousstudies,Neogene 2005;RojayandKaraca,2008).Afterthedevastatingearthquakes

sedimentsoftheKazanregionwereincludedintheBeypazarıBasin. in recent years (Izmit, 1999 and C¸ankırı, 2000), research on the

Aswillbeseenbelow,thisconceptionmostbereconsidered.The tectonics and seismic activities largely focused in the Kazan-

BeypazarıandKazanbasinsform,togetherwiththeKVP,ageomor- Kızılcahamam region (Koc¸yig˘it et al., 2001; Karaca, 2004; Toori,

phological and climatic transition zone between the central and 2005; Seyitog˘lu et al., 2009; Kahraman, 2014). However, a com-

northernregionsofAnatolia(Fig.1).Semi-aridclimate,steppicand monprobleminalmostallstudiesisfrequentlaterallithological

ratherflatlandscapesoftheAnatolianplateau,whichisbetween changes in terrestrial Neogene units in the region and, conse-

thealtitudesof850–1100meters,turnnorthwardtoamountainous quently,dissimilaritiesinlocalstratigraphies(seeEris¸enandÜnlü,

morphologysplitbydeepvalleysandundertheregimeoftherainy 1976;Eroletal.,1980;Türkecanetal.,1991;DuruandAksay,2002;

climateoftheBlackSearegion.Thistransitionzoneisknownwith Lunkkaetal.,2003).Toresolvethiscomplexityneedsmorefossils

itsgreatfloralandfaunalbiodiversity,itsmixedvegetationdom- forahigh-resolutionbiostratigraphyand/orradiometricdating.

inatedbyQuercusandPinus,andalsoothertransitionforesttype

plants(AdıgüzelandVural,1995;UyarandC¸etin,2001).Toprotect 2.2. KazanBasin

and to set forth such a particular ecology, this region became in

1959theSog˘uksuNationalPark.Theregionisrenownedforitsmin- South to the North Anatolian Fault (NAF) and Kızılcahamam

eralwaterssources,coalquarriesandfossilrichbeds(e.g.,Fig.1B, Volcanic Province (KVP), these are several large basins such as

spots1–5)(Ozansoy,1965;Kasaplıgil,1977;Alpagutetal.,1996; Beypazarı,C¸ankırı-C¸orumandPolatli-Haymanabasins,andmore

Rückert-Ülkümenetal.,2002;Forteliusetal.,2003;Hatipog˘luand southernTuzgölüandSivrihisar-Beylikovabasins(Fig.1).Theseare

Türk,2009).Duetotherichnessofitsnaturalresources,theMin- theremnantsoflargemarineforelandbasins,whichwereformed

istryofEnvironmentintendedtodeclaretheKazan-Kızılcahamam in late Cretaceous-Paleogene along the southern margin of the

regionageopark(Kazancıetal.,2007;Kazancı,2012). Pontidecollisionalzone(Görüretal.,1998;DirikandErol,2000;

From the Paleozoic onwards, Anatolia played the role of tec- Karadenizlietal.,2003).Theycomeintocontactinthestudyarea.

tonic crossroad in the geological history of the Old World. The Consequently, Neogene units north of Ankara and in the Kazan

most significant rock samples of this history can be seen in the regionhavebeenmappedaspartoftheBeypazarıNeogeneBasin

northCentralAnatolia,inparticularintheAnkararegion(Fig.1). insomestudiesorpartoftheC¸ankırı-C¸orumBasinintheothers

ThevariousglobaltectonicmodelsontheevolutionofPaleotethys (Türkecanetal.,1991;DuruandAksay,2002;MTA,2002).Inaddi-

andNeotethyshavebeendevelopedovertherocksofthisregion tion,moststudiesdealingwiththegeologyoftheregion,named

(seeTekeli,1981;OkayandGöncüoglu,2004;DilekandThy,2006, stratigraphic units regardless of the sedimentary basin in which

andreferencestherein).Thetermandconceptof“Melange”were theyare(e.g.Ruprecht,1956;Tatlı,1975;Eris¸enandÜnlü,1976;

shaped on the rocks of this region (Bailey and McCallien, 1953; Eroletal.,1980;Göktenetal.,1988;Lunkkaetal.,2003).Thiscon-

Sengör,2003).Therockunitsandstratigraphyincreaseindiver- fusingsituationisprobablyduetolithologicsimilaritiesofNeogene

sityintheareafromPaleozoiconwards,becauseoftheprogressive deposits:MiocenesedimentaryunitsinCentralAnatolianbasins

grouping of several tectonic units during the Mesozoic (Sakarya aremainlylacustrineandfloodplaindeposits,whereasthePliocene

Zone, Istanbul Zone, Taurides, Pontides) on which various sedi- units are of alluvial and fluvial origin. Moreover, not only the

mentary basins (e.g. C¸ankırı-C¸orum, Beypazarı, Polatli-Haymana, lithologiesbutalsofossilcommunitiesdisplaygreatresemblances

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

context.AnnalesdePaléontologie(2017),http://dx.doi.org/10.1016/j.annpal.2017.01.004
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Fig.1. LocationandgeologicalsettingoftheKazan-C¸es¸tepearea.A.BroadtectonicelementsofTurkeyandsurroundings.NAFZ:NorthAnatolianFaultZone;EAFZ:East

AnatolianFaultZone;DSFZ:DeadSeaFaultZone;IAESZ:Izmir-Ankara-ErzincanSutureZone;KVP,CVP,EAVP:Kızılcahamam,Cappadocian,EastAnatolianvolcanicsprovinces,

respectively.NotethatNAFZisaplateboundarypartlyonanoldsuturezone(IAESZ).B.RelationsofKızılcahamamVolcanicProvince(KVP)withfivesignificantTertiary

sedimentarybasinsofCentralAnatolia.NotethattheKazanBasinisintroducedhereforthefirsttime.C.GeneralizedgeologicalmapofnorthwestAnatolia(fromMTA,2002).

Fossilsites:1C¸es¸tepe,2Sinapmammallocalities,3C¸alta,4Pelitc¸iksilicifiedtrees,GüvemandKeseköyplants,insectsandvertebratelocalities,6C¸ankırıBasinmammal

localities,7C¸orakyerler.

LocalisationetcontextegéologiquedelarégiondeKazan-C¸es¸tepe.A.ÉlémentstectoniquesmajeursdeTurquieetdesrégionsvoisines.NAFZ:zonedelafaillenord-anatoliennedu;

EAFZ:zonedelafailleest-anatolienne;DSFZ:zonedelafailledelaMerMorte;IAESZ:zonedesutured’Izmir-Ankara-Erzincan.KVP,CVP,EAVPsontlesprovincesvolcaniquesde

Kızılcahamam,deCappadoceetdel’Anatolieorientale,respectivement.NotezqueNAFZestunelimitedeplaquesituéeenpartiesuruneanciennezonedesuture(IAESZ).B.Relations

deprovincevolcaniquedeKızılcahamam(KVP)aveccinqbassinssédimentairestertiairessignificatifsdel’Anatoliecentrale.NotezquelebassindeKazanestprésentéicipourla

premièrefois.C.Cartegéologiquegénéraliséedunord-ouestdel’Anatolie(MTA,2002).Sitesfossilifères:1C¸es¸tepe,2localitésàmammifèresdeSinap,3C¸alta,4siteàarbressilicifiés

dePelitc¸ik,5siteàplantes,insectesetvertébrésfossilesdeGu¨vemetdeKeseköy,6localitésàmammifèresdubassindeC¸ankırı,7C¸orakyerler.

from one region to another without any kind of endemism etal.,2002).Unliketheotherbasins,theKazanBasinhasrelatively

(Kazancıetal.,1999;Forteliusetal.,2003;Sarac¸,2003).However, thickPliocenedepositsdevelopedinalluvialfanandmeandering

when studied in detail, Neogene deposits in the Kazan region riverdepositionalenvironments.Thesedifferencesinsedimentary

aredifferentintheirarchitecturefromthoseoftheneighbouring depositswiththoseofsimilarageunitsintheneighbouringbasins

basins. The most important difference is the lack of evaporitic indicatethattheKazanareafunctionedasanindependentbasin

deposits in the Kazan Basin, while they form thick sedimentary duringtheNeogene.Thenorthernpartofthisbasinisfilledwith

unitsinotherbasins(C¸ayirhanFmandKirmirFmintheBeypazari the Kizilcahamam volcanics. It is delimited eastward and west-

Basin, Bayindir Fm and Bozkir Fm in the C¸ankırı-C¸orum Basin; wardbythebasementrocks(Permo-Triassic,Cretaceousandearly

Inci et al., 1986; Karadenizli, 1995, 2011; Kaymakc¸ı, 2000; Varol Paleogenemetamorphicandsedimentaryunits).Thelimitsofthis

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

context.AnnalesdePaléontologie(2017),http://dx.doi.org/10.1016/j.annpal.2017.01.004
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according to their own age data (e.g. Lunkka et al., 2003; Toori,

2005; Kahraman, 2014). Consequently, the geological evolution

of the Ankara region during the Neogene appears particularly

complicatedandconfusing.Itisbeyondthescopeofthisstudyto

discussthestratigraphiesgiveninpreviouspublications.Weaim

heretointroducethelocalstratigraphywiththeunitnamesused

byLunkkaetal.(2003)forsedimentaryrocksandbyTürkecanetal.

(1991)andDuruandAksay(2002)forvolcanicrocks(Figs.2and3).

Inthestudyarea,Miocene-Plioceneandyoungerrocksuncon-

formably overlay the older (pre-Neogene) units (Fig. 2). The

youngest unit of the pre-Neogene rocks is the marine middle

Eocene Güvenc¸ Formation, which is formed of Nummulites and

Assilinarichlimestones,marlsandargillaceouslimestones(Fig.3).

Thisunitrepresentsthelatestprogressofmarinetransgressionin

CentralAnatolia(C¸ayrazFmintheHaymanaBasin,OrhaniyeFm

northofAnkara;Göktenetal.,1988;Toori,2005).

Neogene units formed after the Alpine orogeny, which cover

largeareasinCentralAnatolia,aregenerallymuchlessdeformed

than the older units, and they show often lateral and vertical

lithological changes. Lacustrine deposits are common during the

earlyandmiddleMiocene,whilefluvialandalluvialsystemsdom-

inate during the late Miocene and Pliocene. In the basins that

surroundtheKızılcahamamvolcanicprovince(KVP),sedimentary

infillshowsgreatdiscrepancies;forinstancewhileintheC¸ankırı-

C¸orum and Beypazarı basins, late Miocene and early Pliocene

evaporitelakesoccupiedlargeareas,(Fig.1B,C),evaporitedeposits

arenotknownintheNeogenesedimentarysuccessionoftheKazan

Basin.TheNeogeneunitsoftheKazanBasinaresummarizedhere

belowfromthebottomtothetop.

PazarFormation:Thisunitiscommonlyobservedinthestudy

area.ItwasdefinedbyTatlı(1975)aroundPazartown,whichisbuilt

onyoungerdeposits(!),about5kmtothenortheastofC¸es¸tepe

(Fig.3).ThePazarFormationlaterallyinterfingerswiththeKizilc-

ahamamvolcanics(Figs.2and3).Themainlithologiesareclayey

limestone,marlandclaystone,locallychertylimestone,sandstone

andtuff,withoccasionalthincoalseams.Thedominantcolorofthe

unitiswhiteandlightgray.

Locally,dependingontheabundanceandalterationoftuffinput,

white, green or yellow levels can be observed. Its thickness was
Fig.2. StratigraphyofNeogeneunitsintheC¸es¸tepe-Pazararea(modifiedfrom
measurednearKazantobeabout250m(Karaca,2004).Thisfor-
LSturnaktikgara epth aile.,d 2e0s0u3n)i,t aénsdn époogsèintieosnd oafn tshlae rC¸ éegs¸i toenpdee vCe¸erts¸etebprea-tPea lzoacra(limtyo.difiéedeLunkka mationma ybe observ ed as wellin low la ndsand altitud eterr ain,

etal.,2003),etpositiondelalocalitéàvertébrésdeC¸es¸tepe. becauseitisgreatlyfaultedandfoldedbytectonicactivity.Accord-

ingtoitsmainlithologies,thePazarFormationwasformedinlake

andswampenvironmentswithwaterdepthvaryingintimes(Tatlı,
basin,dependingonthepresentdayobservations,aremarkedon
1975).Thelateralequivalentsoftheunitwerenamedinprevious
Fig.1B.Formoreaccurateboundaries,detailedstudiesareneeded.
studies as Akhüyük Fm (Erol et al., 1980), Hanc¸ılı Fm (Duru and
The term “Kazan Basin” was previously used by Toori (2005)
Aksay,2002),C¸eltikc¸iFm(Rojay,2013).Theseauthorssuggestedto
without, however, a clear definition of its characteristics (origin,
dateit,basedonrelativestratigraphyorpollenrecord,asMiocene,
architecture, boundaries). It is out of the purpose of the present
middle-lateMioceneorlateMiocene.Theseformationnamesand
studytoinvolvefurtherinthedefinitionoftheKazanBasinandits
agescannotbeappliedatleastfortheKazanarea.Becauseofthe
detailedgeologicalfeatures.Itsstratigraphyandthepositionofthe
occurrenceofhominoidprimates,thesedimentarydepositsofthe
C¸es¸tepefossillocalityareresumedherebelow.
Kazanareahavebeenintensivelyinvestigatedforstratigraphy,sed-

imentologyandvertebratepaleontology.Theyhavebeenprecisely

2.3. StratigraphyoftheC¸es¸tepe-Kazanarea datedbyfossils,magnetostratigraphyandradiometricdating(Sen,

1991;Alpagutetal.,1996;Forteliusetal.,2003).Thebasalticflow,

The geology of the Kazan-C¸es¸tepe region is shown on Fig. 3. whichseparatesthePazarFormationfromtheoverlayingSinapFor-

Several geologists mapped the geology of the region north of mationisdatedto15.8mybytheK-Armethod(Kappelmanetal.,

Ankara, and they provided various ages for stratigraphic units 2003).ThisbasaltflowisdifferentfromtheÖzlübasaltgivenon

and consequently different tectonic interpretations. Although Fig.3.Alsothemammallocalities24and24AattheInönüdistrict

in almost all geological maps the outcrop limits are similar, the nearSarılarvillage,whicharecorrelatedtothemammalianzone

stratigraphy and unit names are different from one study to MN6(Forteliusetal.,2003)areinthisformation.Allthisshowsthat

another(e.g.Ruprecht,1956;Tatlı,1975;Eris¸enandÜnlü,1976; thePazarFormationwassettledduringtheearlyandearlymiddle

Öngür, 1976; Erol et al., 1980; Türkecan et al., 1991; Duru and Miocene,atleastintheKazanarea.

Aksay,2002;MTA,2002;Karaca,2004;Rojay,2013).Ontheother Kızılcahamam Volcanics: In the study area, Neogene volcanic

hand,thestudiesnotdealingwithmappingusedstratigraphicunit rocks are observed laterally interfingering with the Pazar For-

namesgivenbythepreviousones,butreorganizedthestratigraphy mation. They represent a small part of the southern section of

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

context.AnnalesdePaléontologie(2017),http://dx.doi.org/10.1016/j.annpal.2017.01.004
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Fig.3. DetailedgeologicalmapoftheregionsituatedbetweenthetownsofPazar,KazanandC¸alta.SeeFig.1forfossiliferoussitesintheregion.

CartegéologiquedétailléedelarégionsituéeentrelesbourgsdePazar,KazanetC¸alta.VoirFig.1pourlessitesfossilifèresdelarégion.

ModifiedafterDuruandAksay,2002.

theKizilcahamamVolcanicProvince(KVP).Themainpartofthis earlyPliocene.InseveralbasinsofCentralAnatolia,similardeposits

province is situated further north and spread across an area aremapped,asshownintheMTAmaps,as“undifferenciatedclas-
of 12.500 k m2. It cu lminates at Kö rog˘lu Tepe t o an al titu de of tics ” or Ören cik Form ati on ( Duru and A ks ay, 2002; MTA, 2 002).

2400meters(Figs.1band3).Thisvolcanicprovinceisalsoknown Inindividualstudies,itisnamedasGölbas¸ıFmsouthofAnkara,

in the literature as Körog˘lu volcanics, Galatia Massif and Galatia Etimesgut Fm (Erol, 1980), Memlik Fm (Gökten et al., 1988) and

Volcanics(Türkecanetal.,1991;Topraketal.,1996).Itsmainrock YuvaGroup(Koc¸yig˘itetal.,1988)northofAnkara.Karaca(2004)

typesareandesitic,trachyticandbasalticlavas,rhyolitictuffsand andRojayandKaraca(2008)haveusedthenameofSinapForma-

pyroclastics. There are many volcanic eruptive centers, most of tion,butgavetheageoftheunitasPlio-Quaternarywithouttaking

themaroundKizilcahamamtown,andtheprominentcenterbeing intoconsiderationpaleontologicalandmagnetostratigraphicages

Körog˘luTepe.Accordingtoradiometricdata,thesearetheprod- already available in the literature. In all these studies, the age

uctofthreephasedexplosivevolcaniceventsbetween23-11my ofthisunitisgiveninrelativetermsbasedontherelativeposi-

(Türkecan et al., 1991; Wilson et al., 1997; Tankut et al., 1998). tionsimilarlithologiesinotherbasins.Forteliusetal.(2003)and

Duringthecalmperiods,lakesoccupiedlargeareas,andterrestrial Lunkkaetal.(2003)carriedoutin1990sdetailedsedimentological,

sedimentarydeposits,whichalsolocallyincludecoalformations, stratigraphicandinparticularpaleontologicalinvestigationsinthe

developed.Thesedepositsalsoincludeseveralfossiliferouslocal- KazanBasin(Fig.3),mainlyconcentratedaroundthetypesectionof

ities, the best known of which are Güvem-Keseköy with leave theSinapFormation.Lunkkaetal.(2003)distinguishedfivesucces-

and vertebrate fossils and Bayındır Pelitc¸ik silicified wood forest sivemembers(fromthebottomtothetopBeycedere,Yellidoruk,
(loca lities4and 5onF ig.1 B)(Kasap lıgil, 19 78;Rück ert-Ülk ümen I˙g˘be k,Kavakde reand C¸al tamem ber s),c ove ringaninte rvaloftime

etal.,2002;AtabeyandSarac¸,2004;Lopez-Anton˜anzasetal.,2004; fromthemiddleMiocenetotheearlyPliocene(Fig.2).Themam-

Hatipog˘luandTürk,2009;Kazancı,2012).Themajorityofvolcanic malian faunas from the first four members have been subject of

rocks in the study area are basaltic lava and coarse pyroclastics detailed studies published in a volume edited by Fortelius et al.

of the Ilıcadere volcanics (Türkecan et al., 1991) that are mostly (2003),whilethefaunafromC¸altawaspublishedinavolumeedited

derivedfromvolcaniccentersnearKızılcahamam(Fig.3). bySen(1998a,1998b).TheoutcropsseenaroundC¸es¸tepe,Pazar

SinapFormation:Itcoversthelowlandsofthestudyareainthe andKurtbogazilocalitiesallbelongtotheC¸altaMember.Itsthick-

formofthreepatchesaroundKazan,C¸altaandC¸es¸tepe-Pazar(Pls nessnearC¸es¸tepeisabout45m,anditiscomposedoffloodplain

onFig.3,PlonFig.1).Atypicalerosionalsurfaceisdevelopedon deposits(mudstones)cutoffbystreamchannelsediments(loose

itsoutcrops.Thisformationiseasilydistinguishedfromtheunder- conglomerates,sandstones).AroundtheC¸es¸tepevillage,wherethe

lyingPazarFormationbyitspoorlyconsolidatedsedimentswith vertebratefossilsarerecorded,fossiliferoussedimentscorrelates

yellowish,reddish,mottledgray,browncolors.Itisalmosttotally tothemiddle-upperhorizonsoftheC¸altaMember.Mudstonesand

composed of lithologies of alluvial and fluvial sediments domi- gravellychanneldepositsaredominant(Fig.4A–D).Sedimentsare

natedbylooseconglomerates,sandstonesandmudstones.Locally not well consolidated, and consequently deeply cut by seasonal

itincludeslacustrinesedimentsthatformthinbands.Thisforma- streamsallowingthedevelopmentofarecentfluvialnetworkon

tionisnotmuchaffectedbythetectonics,andthusthebeddingis theC¸altaMember.

almosthorizontal(RojayandKaraca,2008).ItcoversPazarForma-

tionandolderunitswithangularunconformity(Fig.2).Although 3. Materialandmethods

itsusualthicknessisbetween40–100m,Karaca(2004)measured

asectionof150mnorthwesttoKazan. ThevertebrateremainsfromC¸es¸tepehavebeenfoundduring

ThisunitwasdefinedbyTatlı(1975)alongtheslopesofSinap thediggingoperationsforthefoundationofaprivatehouseatthe

Tepeatsome4kmnorthwesttoKazan,withagivenagerangeas villageofC¸es¸tepe.AdemBüyükköprü,ageologistwhoworksatthe

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

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Fig.4. Close-upviewtothefossiliferousPliocenedepositsatC¸es¸tepe.A.Floodplainmudstones(withachannel)unconformablycoveredbyQuaternaryterracedeposits.

ThelatterformsalsoamorphologicalsurfacethroughouttheKazanbasin.Thewhitishcolorofthesedimentisduetofinepyroclasticparticles.B.Mudstonesofanalluvial

floodplain(=distalfan)overlinedbyfluvialsandydeposits.ThisisacommonsedimentpatternofthePliocenesequencesintheKazanBasin.C.Alternationofalluvialand

fluvialdeposits.ThisisatypicalfeatureofsedimentsoftheSinapFormation.D.Vertebraeofmonitorlizard(Varanussp.)atC¸es¸tepeinbluishmudstonespossiblyrepresenting

ashallowpondormarshinafluvialsetting.

VuerapprochéedesdépôtsfossilifèresduPliocèneàC¸es¸tepe.A.Bouedeplained’inondation(avecunchenal)couverteendiscordancepardesterrassesquaternaires.Cesderniers

formentégalementunesurfacemorphologiquedanstoutlebassindeKazan.Lacouleurblanchâtredusédimentestdueàdefinesparticulespyroclastiques.B.Lesmudstonesdeplaine

alluviale(=éventaildistal)surmontéededépôtsfluviauxsableux.CetypedesuccessionsédimentaireestcommundanslesséquencespliocènesdubassindeKazan.C.Alternancedes

dépôtsalluviauxetfluviaux.CetypededépôtestcaractéristiquepourlaFormationdeSinap.D.Vertèbresd’unvaran(Varanussp.)àC¸es¸tepedansdesargilesbleutéesprobablement

déposéesdansunétangpeuprofondouunmaraisdansunmilieufluvial.

DepartmentofWaterSupplyandSewerageofthecityofAnkara, Forthedescriptionofochotonidteeth,weusedtheterminology

notedthepresenceofbonesonthesection(Fig.4D).Hestopped ofLópez-Martínez(1989).Uppercheekteethareabbreviatedwith

diggingandwarnedonthisdiscoveryoneofus(N.Kazanci)atthe capital letters and lower cheek teeth with lower case letters. All

University of Ankara. A team from this university went the next measurementsareinmillimeter.

day to C¸es¸tepe and collected all visible bones and some matrix

at the place of discovery. The matrix was sieved on a mesh of 4. Systematicstudy
1m mt oreco ver fragments not easilyvi sible .Thege ogr ap hiccoo r-
d inate s of the fi nding plac e ar e N 4 0◦ 19(cid:4) 3 0.8 9(cid:4)(cid:4) and E 32 ◦ 41(cid:4) ClassReptiliaLaurenti,1768
21.85(cid:4)(cid:4). Order Squama taOppel ,1811

Gelogists who collected this material at C¸es¸tepe are not SuperfamilyLacertoideaOppel,1811

familiarwiththetechniquesofdiggingandcollectingfossilverte- FamilyLacertidaeOppel,1811
brates.T heyh ave collectedth ef ossilsw itho utconsolid ation .Fresh Genus LacertaLin naeus, 1758

fractures on bones, the presence of numerous small fragments

belongingtoanochotonidskull,fragmentarystateoflizardverte- cf.Lacertasp.

braethatapparentlybelongedtothesameindividualshowthatthe Material:onefragmentaryrightdentary(CT-10).

materialwasinitiallyconsistedofmorecompletespecimens,but Description: The incomplete right dentary CT-10 (Fig. 5A, B)

damagedduringexcavationandsalvageoperations. preserves the anterior portion, 8mm long, up to the eighteenth

ThematerialfromC¸es¸tepeincludestheremainsofanochotonid tooth position. Teeth are (at least partially) preserved in the

lagomorphandseverallizards.Lizardsarerepresentedby88spec- positions from two to seven, nine, 11, 13, 15 and 16; they are

imensreferredtothreetaxa:cf.Lacertasp.,non-AnguisAnguinae pleurodont,longandcylindrical.Theanteriorteethareanterodor-

indet.andVaranussp.Thenumerousfragmentedvertebraeprob- sallydirectedandshowasinglecusp.Theotherteetharedorsally

ably belonged to at least partially complete skeleton of monitor directed and the well-preserved crown in the ninth position is

lizardasseenonFig.4D.Theochotonidisrepresentedbythreefrag- clearlybicuspid,withaverysmallanteriorcuspletseparatedbya

mentarylowerjaws,skullfragmentsandoneuppercheektooth. shallowlingualgroovefromthemaincusp.Otherteeth,astheone

We tried to fix bone fragments to complete some specimens as preservedintheeleventhposition,showalingualgrooveprobably

much as possible, without great success due to the loss of con- indicatingtheformerpresenceofananteriorcusplet.Thedental

tactbetweenthefragments.AllthismaterialishousedatAnkara crestisdorsallycurvedanditsmedialedgeisrounded.TheMeckel’s

University,departmentofEngineeringGeology. grooveisfullyopen,reachingtheanteriortipofthedentary,and

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

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Fig.5. A,B.cf.Lacertasp.,fragmentaryrightdentaryCT-10inlateralandmedialviews.C.Non-AnguisAnguinaeindet.,poorlypreservedosteodermCT-11inexternalview.

D–F.Varanussp.,trunkvertebraCT-12indorsal,ventral,andleftlateralviews.Scalebarsequal2mminA–Cand10mminD–F.

A,B.cf.Lacertasp.,fragmentdedentairedroit(CT-10)envuelatéraleetmédiale.C.Non-AnguisAnguinaeindét.,ostéodermefragmentaire(CT-11)envueexterne.D–F.Varanussp.,

vertèbredutronc(CT-12)envuedorsale,ventraleetlatéralegauche.Leséchellescorrespondentà2mmpourA–Cetà10mmpourD–F.

widensposteriorly.Inlateralview,thedentarydeepensbackward Material:1osteoderm(CT-11).

and hosts six mental foramina aligned to the dorsal edge of the Description: The preserved portion of the osteoderm CT-11

element. (Fig.5C)is5.0mmwideand5.1mmlong.Thelateralandposterior

Remarks: The characters described above, the pleurodont edgesareincomplete.Theexternalsurfaceisaltered,butthefol-

bicuspiddentitionandtheopenMeckel’scanalinparticular,fitwell lowingcharacterscanbedetected:presenceofananteriorsmooth

withthoseoflacertidlizards(amongothersArnold,1973,1989; stripposteriorlyfollowedbyamuchlargerandvermiculatedarea

BarbadilloandSanz,1983;BarahonaandBarbadillo,1997;Arnold dividedintwobyalongitudinalridge.Theinternalsurfacedoes

et al., 2007). The size of the only available element suggests the notpreserveanyrelevantcharacter(i.e.thepresenceofforamina

presenceofamemberofLacertathataccordingtoUetzetal.(2016) cannotbeascertained).

iscurrentlyrepresentedinAnatoliabythefollowingspecies:Lac- Remarks: Isolated osteoderms are very common in the fossil

erta agilis Linnaeus, 1758, Lacerta media Lantz and Cyrén, 1920, record(forexample,morethan8000inthelateMioceneofMon-

Lacerta pamphylica Schmidtler, 1975, Lacerta strigata Eichwald, cuccoTorinese–Italy;Colomberoetal.,2017)probablybecause

1831,LacertatrilineataBedriaga,1886,andLacertaviridis(Laurenti, they are massive bony structures that are very numerous in a

1768).However,sincethecomparativeosteologyofthesespecies single lizard individual and they are also easy to detect because

is currently unknown and, moreover, it cannot be excluded that of their characteristic shape and relatively large size. However,

thesinglefragmentarylacertidelementcurrentlyavailable(CT-10) eveniftheskullandlowerjawosteologyofthewesternPalaearc-

belongstoasubadultrepresentativeofthelargergenusTimonor ticanguineswascarefullydescribed(amongothers:Rocˇek,1980;

eventoanextincttaxon,theidentificationislimitedatgenuslevel, Klembaraetal.,2010,2014;Klembara,2012),littleattentionhas

withtheadditionofthequalifiercf.indicatingclosemorphological beenpaidtothecomparativemorphologyoftheosteodermsand

similaritytoLacerta. thereforeonlyAnguiscanbeeasilyandreliablydetected.Asalready

Lacertidlizardswerereportedfromabout10Turkishlocalities remarked by Colombero et al. (2014), the osteoderms coming

whoseagerangesfromthelateOligocenetotheMiddlePleistocene from a certain site are usually referred to the taxon identified

(Böhme and Ilg, 2003). Some of them have been even referred onthebasisofcranialcharacters,andwhentwotaxaarerecog-

tothegenusLacerta,butonlythosefromthemiddlePleistocene nizedinasinglesiteonthebasisofcranialelementsithasbeen

of Emirkaya (Venczel and Sen, 1994) have been described and suggested not to refer the osteoderms to them (Augé and Rage,

figured. No other lacertid genera than Lacerta were reported for 2000).

Turkeydespitethefactthatitcurrentlyhosts13genera(Uetzetal., Anguid lizards were reported in about 15 Turkish localities

2016).Inthesamebroadgeographicarea,agreenlizard,Lacerta (Böhme and Ilg, 2003), but with the exception of the materials

gr.L.viridis,hasbeenrecentlyreportedfromtheEarlyPleistocene referred to Pseudopus from the early Pliocene of C¸alta (Rage and

ofDmanisiinGeorgia(Blainetal.,2014)andLacertasp.,Ophisops Sen, 1976; Sen and Rage, 1979) and the middle Pleistocene of

elegansandLacertidaeindet.fromtheMiddlePleistoceneofAzokh Emirkaya(VenczelandSen,1994),alltheotherswereidentified

1CaveinNagornoKarabakhregion(Blain,2016). atfamilylevel.Pseudopusapodusistheonlynon-Anguisanguine

currentlyinhabitingAnatolia(Uetzetal.,2016)andwasrecently

reported from the Middle Pleistocene of Azokh 1 Cave (Nagorno
SuperfamilyAnguioideaGray,1825
Karabakhregion;Blain,2016).
FamilyAnguidaeGray,1825

SubfamilyAnguinaeGray,1825

SuperfamilyVaranoideaCamp,1923

non-AnguisAnguinaeindet. FamilyVaranidaeGray,1827

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GenusVaranusMerrem,1820 Table1
Measu rementsofloweranduppercheekteethofOchotonamediterranensisfrom

C¸es¸tepe,Turkey.
Varanussp.
Dimensionsdesdentsjugalesd’OchotonamediterranensisdeC¸es¸tepe,Turquie.
Material: two trunk vertebrae (CT-12 and 13); 20 fragments
referable to at le ast 15 trunk ver tebrae (CT-1 4); 1 fr agmentary Tooth n Length Widthtrigonid Widthtalonid

sacralvertebra(CT-15);15fragmentarycaudalvertebrae(CT-16); Range X Range X Range X

31 vertebral fragments (CT-17); 1 fragmentary right scapuloco-
p3 1 1.36 1.25
racoid (CT-18); 1 fragmentary right ischium (CT-19); 1 distal
p4 2 1.38–1.47 1.43 1.33–1.43 1.38 1.48–1.54 1.51
epiphy sisofa(? )tib ia(CT-20);1 righta stragaloc alcaneum (C T-21); m1 3 1.48–1.51 1.50 1.49–1.57 1.52 1.56–1.61 1.58

1tarsal(CT-22);5metapodials(CT-23);7phalanges(CT-24). m2 3 1.30–1.53 1.45 1.42–1.57 1.51 1.49–1.55 1.52
Desc ription: D espitenotbe ingcomp le te,becaus etheanterior P4 1 1.08 1.64 1.97

edge of the neural arch, the tip of the neural spine, the dorsal n:numberofspecimens;X:meanvalues.

portionofthecotyleandthecondylearepartiallydamaged,the

trunkvertebraCT-12(Fig.5D–F)isthemostinformativeelement OrderLagomorphaBrandt,1885

of this taxon. The preserved portion of the anterior area of the FamilyOchotonidaeThomas,1897
ne ural archisc onc ave.Thepre zygapop hy sesa remarke dlyt ilte din Genus OchotonaLin k,1795

dorsolateraldirection.Thesynapophysesareratherglobularand

noticeablyprotrudinglaterally,butwithanarrowconnectionwith OchotonamediterranensisSuataAlpaslan,2009

the lateral edge of the cotyle. The neural canal is proportionally Material:IsolatedrightP4(CT-1);rightmandiblefragmentwith

narrow and roundish. The cotyle and the condyle are markedly p3-m2(CT-2);rightmandiblefragmentwithdetachedincisor,alve-

compresseddorsoventrally,withaventraledgeverymuchmore olusofp3andp4-m2(CT-3);rightmandiblefragmentwithm1-m2

posteriorlyplacedthanthedorsaledge.Thelateralsidesofthever- (CT-4),fragmentoframusofarightmandible(CT-5).Thematerial

tebraandpartofthoseoftheneuralarcharedistinctlystriated.The alsoincludesseveralsmallfragmentsthatapparentlybelongedto

centrum,11.7mmlong,showslateraledgesdistinctlyconverging askull.Wetriedtofixthemtorebuildthisskull,withoutsuccess.

posteriorlyandwithasignificantprecondylarconstriction. Theanatomicalpositionofmostfragmentsisnotrecognizable.Only

Caudalvertebraearecharacterizedbythefollowingcharacters: onepairofoccipitalcondylestogetherandpartsofoccipitalbones

cotyleandcondylesimilartothosedescribedforthetrunkverte- arereliablyreferabletothisochotonid.Thematerialishomogenous

brae; centrum anteroposteriorly elongated; lateral apophyses anddoesnotcontainfossilsthatmaybelongtoanyothertaxon.

dorsoventrally flattened and variably broadened; pedestals for Measurements:SeeTable1.

chevronbonespresentandwell-defined;precondylarconstriction. Description:Puttingtogethertheavailablefragments,itispos-

Thelargestcaudalvertebrawithameasurablecentrum(CT-16a) sibletorestorethemandible(Figs.6and7).Themissingpartsare

is9.8mmlong. theposteriorpartofthebody,partoftheangularprocessandm3.

Therestoftheremainsareveryfragmentaryand/ornottaxo- Inlateralview,thebodyisdeepestbelowp4(5.20mm),whilethe

nomicallydiagnostic(astheastragalocalcaneumCT-21)becauseof ventral margin of the bone shows a clear swelling forward and

theabsenceofcomparativestudiesontheirmorphology. backward.Thetipoftheincisorreachestheocclusallevelofcheek

Remarks: The size and general morphology of the vertebrae, teeth.Thelengthofthediastemais4.58mmonthemostcomplete

thedorsoventrallycompressedcotyleandcondylewiththeirven- specimen(CT-2),anditisgentlyconcave.Thep3-m3lengthcan

traledgemuchposteriorthanthedorsalone,andtheprecondylar beestimatedasabout7mm.Belowthep4,thereisastrongtaurus

constrictioninparticular(anautoapomorphyofVaranus,seeSmith mandibulae,whichisaroundbonethickeningfortheattachment

et al., 2008, and literature therein), clearly support the identifi- ofmassetermuscles.Thelateralsurfaceofthebodyhasnumerous

cation of a monitor lizard of the genus Varanus. However, their tinyforamina,mainlygroupedbelowp3andm1.Theanteriormen-

morphologyisnotcongruentwiththatofVaranusgriseus(Daudin, talforamenissituatedbelowthep3(CT-3)orp3-p4limit(CT-2).

1803),theonlyspeciescurrentlyinhabitingAnatolia(withasin- Theposteriormentalforamenissituatedabout3.5mmbelowthe

gle taxon/locality datum in southeastern Turkey; Sindaco and m2.

Jeremcenko,2008).Thedistinctlyconverginglateraledgesofthe Onthefragmentoframus,thearticularprocesshasanarticular

centrum(inventralview)andthereforethemarkedprecondylar surfacethatcoverstheheadofthisprocess.Dorsallyitisteardrop

constriction of the vertebra CT-12 are in contrast with the mor- shaped.Thecoronoidprocessisnotpreserved.Theangularprocess

phologyofV.griseus,butreminiscentofageneralizedmorphology ispartlypreservedanditisapparentlyquitestrong.

of many other species with at least a relatively large size (see Inmedialview,thearticularsurfaceofthesymphysisextends

fig. 3 in Delfino et al., 2013). Such morphology was reported in on 2.35mm below the diastema. The medial face of the body is

theMediterraneanareaforboththelateMioceneextinctVaranus concavebetweenthealveolarmarginandtheprotuberanceofthe

marathonensis Weithofer, 1888 and Varanus amnhophilis Conrad lowerincisor.Severalsmallforaminaarescatteredonthisface.

etal.,2012. In ventral view, the mylohyoid line starts where the lower

Varanus was reported in four Turkish localities (Böhme and incisors ends and it extends backward along the medial side till

Ilg, 2003) ranging in age from the middle Miocene to the early belowm2.

Pliocene,andisolatedtrunkvertebraefromtheearlyPlioceneof A complete lower incisor measures 12.3mm in length. It is

C¸alta (Rage and Sen, 1976) with a morphology similar to those gently curved. The enamel only covers the mesial face (width

fromC¸es¸tepewereactuallyreferredtoV.marathonensis.However, 1.51mm).Thesectionisalmosttriangular.Theanterioredgebears

besidesdoubtsonthevalidityofthelatterspecies,thevertebral twowearfacets,theonedorsalon2.71mmandtheothermedial

morphology is not diagnostic at species level and therefore the on2.50mm,bothseparatedbyarightangle.Insidethemandible,

materialheredescribedissimplyreferredatgenuslevel.Sinceall thedistalendoftheincisorreachesthelevelofp4-m1boundary,

fossil remains from C¸es¸tepe could belong to a single individual anditislinguallyrootednexttothepremolars.

(becauseofthecongruentsizeandabsenceofduplications)allthe From the cheek teeth, the p3 is slightly longer than wide.

restofthematerialisreferredtothesametaxon. Theanteroconidhasabadlydefinedtrefoilocclusaloutlinewith

anterolateral and anteromedial depressions without cement. Its

MammaliaLinnaeus,1758 length and width are similar (0.56mm). The anteroconid is con-

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nectedtotheposteriorpartbyanarrowisthmus.Theparaflexid fromthelabialmargin.Theenamelisthickonthemesialmarginof

isclearlyshorterthantheprotoflexid,andbothareobliqueback- lophsbutthinontheirdistalmargin;itlacksonthelabialmargin.

ward.Thehypoflexidisshallow,V-shapedandrathertransverse. Theshaftismoderatelycurvedlabiallyandwrinkledmesially.

Allthreeflexidsarefilledwithcement.Themesoflexidislacking. Remarks:TheC¸es¸tepeochotonidclearlybelongstothegenus

The enamel is thick around the protoconid, in front of the para- Ochotona in having small-sized cheek teeth, p3 with a simple

conid,andalongtheposteriormarginofthehypoconid.Enamelis occlusalpattern,twolabialandonelingualflexids,theanteroconid

moderatelythickandratherhomogeneousontheanteroconid. ofp3withoutflexidandconnectedtotheposteriorcomplexbya

Thep4,m1andm2arebilophodontasusualinallochotonids. narrowisthmus.

Theanteriorlophisdiamondshaped,whiletheposteriorlophis Among the most similar ochotonids, the genus Ochotonoides

rathercrescentshaped.Theposteriorlophisalittlelargerthanthe TeilharddeChardinandYoung,1931isdifferentinbeingofvery

anterior loph in p4 and m1, but slightly narrower or equivalent largesize,andinhavinglargeanteroconidonp3withfoldedand

inm2.Theenamelisthickontheposteriormarginofbothlophs, cementedflexids.ThegenusOchotonomaSen,1998a,1998bisalso

whichcorrespondstothemainshearingblades,andratherthinon different in being larger in size, and, in particular, in having p3

theanteriormargins;inanycase,itisneverinterrupted. with a large anteroconid and a short talonid; its anteroconid is

AcompleteisolateduppercheektoothisdeterminedasrightP4 penetratedbylabialand/orlingualflexidsfilledwithcement;the

becauseofitsanteriorlophclearlyshorterandnarrowerthanthe protoflexidandparaflexidarelarger,deeperandstronglydirected

posteriorloph(1.64versus1.97mm).Thehypoflexusendsquitefar backward;thehypoflexidisnarrow,andthelophsonp4-m2are

compressedmesio-distally.PliolagomysAgadjanianandErbajeva,

1983 is another ochotonid known from the Pliocene and early

PleistoceneofEasternEuropeandWesternAsia.Thethreespecies

referredtothisgenusarelargerinsizethantheC¸es¸tepespecimens,

their p3 have anteroconid enamel more or less folded, the isth-

musbetweentheanteroconidandposteroconidislabiallyshifted

because the paraflexid is twice deeper than the protoflexid, and

hypoflexidnarrower.

ThegenusOchotonahasalargedistributioninAsiaandwest-

ern North America with 28 extant species generally recognized

based on morphological as well as molecular studies (Hoffmann

and Smith, 2005; Ge et al., 2013; Lissovsky, 2014). Some of the

extant species are also known as fossil during the Pleistocene,

e.g.,Ochotonapusilla(Pallas,1769),whichhasalargedistribution

inEurasiaduringthePleistocene(Malatesta,1951;Malez,1969;

Erbajevaetal.,2001).Inadditiontotheextantspecies,Erbajeva

etal.(2015)recognized44extinctspeciesofOchotonainEuropeand

inparticularAsia.Aswewilldiscussherebelow,thequestionofthe

synonymyofsomespeciesisanongoingdebateamongthelago-

morphspecialists,whichisoutofthepurposehere.Actually,such

aquestioncannotbesolvedwithoutacompleterevisionofavail-

ablecollections,andperhapsmoreabundantandbetterpreserved

specimens.
During the last decade some effort was invested for a better

descriptionoftheEuropeanandsomeAsianfossilspecies(seein

particularErbajevaetal.,2001;Sen,2003;Fostowicz-Frelik,2008;
Fostowicz -Frelikan dF reli k,201 0;Cˇe rmák, 2010,2016;Cˇermá kand

Rekovets, 2010; Delinschi, 2014) and for morphological cladistic

analysisofmostextinctandsomeextantspecies(Fostowicz-Frelik

etal.,2010;Geetal.,2013).Thesestudiesprovidereliablecriteria

forcomparingtheC¸es¸tepeochotonidwithothersforitssystematic

assignment.
Ochotona sp. cf. O. lagreli Schlosser, 1924 from late middle

MiocenelocalityofDamiaoinInnerMongolia(Zhangetal.,2012)

is apparently the oldest record referred to the genus. Ochotona

ozansoyi Sen, 2003 from the late Vallesian of Turkey (dated to

9.4Ma)andOchotonakalfense(Lungu,1981)fromthelateValle-

sianofMoldova(ca.9–9.5Ma)evidencethatthisgenus,probably

originatedinEastAsia,alreadyreachedtheeasternMediterranean-
southeaste rn Eur ope i n late M iocene ( Sen , 2003; Cˇermák, 2016).

O.ozansoyiwasrecoveredinTurkeyinthreelocalities(SinapLoc.

84,Inönü1andKüc¸ükc¸ekmece)(Sen,2003,2016),andO.kalfense

Fig.6. OchotonamediterranensisfromC¸es¸tepe,earlyPlioceneofKazanBasin.A.Right atKalfa(Moldova),allcorrelatedtothelateVallesian.Bothspecies
man di ble(CT-2) withp3-m2inla bialv i ew .B.Sa mesp ecimen in lingua lview. C. Right ar elarg erinsizeco mp aredtoth eC ¸es¸t epe specimens ,hav elarger

mandiblefragment(CT-3)withthealveolusofp3andp4-m2inlabialview.Scale andsimple-shapedanteroconidandshorterposteriorsegmenton
bar equals 5 mm. the p3, and their l ower (upper ) ch eek teet h are cur ved labia lly
Ochotona mediterranensis de C¸es¸tepe, Pliocène inférieur du bassin de Kazan.
(lingually),whileinPlio-PleistocenerepresentativesofOchotona,
CA.. MFraangdmibeunlted deromitaen (dCiTb-u2l)e advreoci pte3-(mCT2- 3en) vauveec lalb’ailavléeo. Ble. Mdeêmp3e septépc4im-men2 eenn vvuuee linlagbuiaallee.. thelowerc heekt ee thbecomerathe rstrait,andthe cu rvatureof

L’échellecorrespondà5mm. upper cheek teeth decrease. We consider the genus Proochotona

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

context.AnnalesdePaléontologie(2017),http://dx.doi.org/10.1016/j.annpal.2017.01.004
G Model
ARTICLE IN PRESS
ANNPAL-2292; No.ofPages15

10 S.Senetal./AnnalesdePaléontologiexxx(2017)xxx–xxx

Fig.7. OchotonamediterranensisfromC¸es¸tepe,earlyPlioceneofKazanBasin.A.RightP4(CT-4)inocclusalview.B.Rightlowertoothrow(CT-3)withthealveolusofp3and

p4-m2inocclusalview.C.Rightlowertoothrow(CT-2)withp3-m2inocclusalview.D.RightlowerjawreconstructedusingallavailablefragmentsfromC¸es¸tepe,inlabial

view.Thegrayedpartsaremissing.Oncheekteeth,blackenedlinesrepresentenamel;white,dentine;dottedareasarecementinfill.Scalebarsequal1mminA–Cand5mm

inD.

OchotonamediterranensisdeC¸es¸tepe,PliocèneinférieurdubassindeKazan.A.P4droit(CT-4)envueocclusale.B.Sériedentaireinférieuredroite(CT-3)avecl’alvéoledep3et

p4-m2envueocclusale.C.Sériedentaireinférieuredroite(CT-2)avecp3-m2envueocclusale.D.Mâchoireinférieuredroitereconstruiteenutilisanttouslesfragmentsdisponibles,

envuelabiale.Lespartiesgriséessontmanquantes.Surlesdents,leslignesnoirciesreprésententl’émail;lespartiesblanchesindiquentladentineetleszonesenpointilléssont

rempliesdecément.Leséchellescorrespondentà1mmpourA–Cetà5mmpourD.

Khomenko,1914asajuniorsynonymofOchotona(seediscussion near Konya (Montuire et al., 1994; Santel and Von Koenigswald,
inSen,2003 ;Cˇerm á k, 2016). OtherOch ot onaspecie sinE uropeand 1998 ;seeal soErbajeva et al., 2001), andCˇe rmák eta l.(2006)men-

TurkeyareallfromPlio-Pleistocenelocalities. tioneditspossibleoccurrenceinGüzyurduCavenearGümüshane.

Thesizeisusedasanimportantcriteriontodistinguishextinct O. pusilla is a small-sized species, characterized by a very small

species of Ochotona. Erbajeva (1988, 1994) grouped Ochotona anteroconid,andreducedparaflexidandprotoflexidonthep3.Itis

species as small, medium and large-sized species. Later on, hardlycomparedwiththeC¸es¸tepeochotonid.

Fostowicz-Frelik et al. (2010) used this criterion in their phy- DuringthePlioceneandearlyPleistoceneinEurope,O.polonica

logenetic analysis. They defined as small the species having is the best-documented species in several localities of Poland.

p3-m3toothrowlengthlowerthan8mm,medium-sizedforlength Fostowicz-Frelik(2008)rightlyincludedinthisspeciesOchotona

between 8.1–9.9mm, and large-sized the species having p3-m3 dehmi from the latest Pliocene locality of Schernfeld (Germany)

longer than 10mm. In this classification, the C¸es¸tepe ochotonid and O. valerotae from Les Valerots (early Pleistocene, France).

ranksamongthesmall-sizedspecies(p3-m3lengthestimatedto She demonstrated that these two species are similar in size to

about 7mm), together with O. bazarovi Erbajeva, 1988, O. dehmi O.polonica,andtheirp3iselongated,theanteroconidissimpleand

Erbajeva,2005,O.mediterranensisSuataAlpaslan,2009,O.minor tapersanteriorlyasinO.polonica.Indeed,afewdifferencesthatthe

Bohlin,1942,O.polonicaSych,1990,O.pusilla,O.valerotaeErbajeva authorsshowedtodistinguishtheseformsfromO.polonicaarein

etal.,2001,O.zabiensisFostowicz-Frelik,2008. therangeofindividualvariation.Themeasurementsofmandible

Among these species, the Central Asian step pika O. pusilla is andcheekteethofO.polonicaarerathersimilartothatofC¸es¸tepe.

quite common in European Plio-Pleistocene localities. Malatesta However, O. polonica is different from the C¸es¸tepe ochotonid in

(1951),Malez(1969)andErbajevaetal.(2001)providedalarge havingmandiblewithflatventralmargin(curvedatC¸es¸tepe),p3

accountofitsoccurrencesinEuropeandAsia.Lastly,Fostowicz- elongated, and its anteroconid elongated and triangular without

FrelikandFrelik(2010)reportedonitsoccurrenceinPolandsince anygroovesandwithasharpapex(Fostowicz-Frelik,2008).The

the latest Pliocene (locality of Kielniki 3), almost coeval with its proportionsoftheanteroconidandposteroconidonthep3arealso

firstoccurrenceinKazakhstan(Aktogai;Erbajeva,1988).InTurkey, different.Insummary,despitethesimilarityinsize,themorphol-

this species was found in two middle-late Pleistocene localities ogyofmandibleandlowercheekteethofO.polonicaareclearly

atYarimburgazCavenearIstanbul,andEmirkaya-2fissurefilling differentfromthatofC¸es¸tepe.

Pleasecitethisarticleinpressas:Sen,S.,etal.,C¸es¸tepe,anewearlyPliocenevertebratelocalityinCentralAnatoliaanditsstratigraphic

context.AnnalesdePaléontologie(2017),http://dx.doi.org/10.1016/j.annpal.2017.01.004