Discovery of an Upper Miocene Vertebrate fauna near Tizi N'Tadderht, Skoura, Ouarzazate Basin (Central High Atlas, Morocco)

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General palaeontology, systematics and evolution (Vertebrate palaeontology)

Discovery of an Upper Miocene Vertebrate fauna near Tizi N’Tadderht, Skoura, Ouarzazate Basin (Central High Atlas, Morocco)

Découverte d’une faune de vertébrés dans le Miocène supérieur de Tizi N’Tadderht, Skoura, Bassin d’Ouarzazate (Haut Atlas Central, Maroc)

Samir Zouhri

^a,∗

, Denis Geraads

^b

, Siham El Boughabi

^a

, Abdelghani El Harﬁ

^c

aLaboratoiredegéosciences,facultédessciences,universitéHassanII-Casablanca,Km8,routed’ElJadida,BP5366Maârif,20100Casablanca,Morocco

bCNRS,UPR2147,44,ruedel’Amiral-Mouchez,75014Paris,France

cLaboratoiredegéologieappliquéeetgéo-environnement,facultédessciences,universitéIbnouZohr,BP28/S,Agadir,Morocco

a rt i c l e i n f o

Articlehistory:

Received24August2011

Acceptedafterrevision9January2012 Availableonline4July2012 PresentedbyPhilippeTaquet

Keywords:

Vertebrates UpperMiocene Ouarzazatebasin TiziN’Tadderht Morocco

a b s t r a c t

ThediscoveryofUpperMiocenevertebratesatTiziN’TadderhtintheOuarzazatebasin (Morocco)helpstoﬁllagapinourknowledgeofNeogenefaunasinNorthAfrica.Thenew faunaincludesanostrichcf.Struthiosp,aturtlecf.Centrochelyssp.,Crocodyluscf.niloticus,andarelativelydiversefaunaoflargemammals.Themammalassemblageprobably includesthreehipparionspecies,includingaverysmallformnotpreviouslyreportedfrom Africa,aff.Cremohipparionperiafricanum,twospeciesofrhinoceroscf.Ceratotheriumsp.

andaff.Chilotheriumsp.,aProboscideancf.Tetralophodonsp.,alargememberoftheGiraf- ﬁdaesimilarto“Palaeotragus”germainiandtwobovidsofwhichoneislikelyrelatedto Prostrepsiceros,whiletheotherisanewmedium-sizedantelopewithspiralhorns,cer- tainlyarepresentativeoftheCaprinae,agroupthatisrareinAfrica.AlateMioceneage, correspondingtotheEuropeanTurolianMammalage,ismostlikelyforthisfauna.

Motsclés: Vertébrés Miocènesupérieur Bassind’Ouarzazate TiziN’Tadderht Maroc

r é s um é

LadécouvertedevertébrésduMiocènesupérieuràTiziN’Tadderht,danslebassinde OuarzazateauMaroc,contribueàcomblerunelacunedansladocumentationfaunique duNéogèned’AfriqueduNord.Cettefaunecomportecf.Struthiosp,cf.Centrochelys,un crocodiledelalignéedeCrocodylusniloticus,ainsiqu’unefaunerelativementdiversiﬁée degrandsmammifères.L’assemblagemammaliencomprendprobablementtroisespèces d’HipparionsdontnotammentuneformedetrèspetitetaillejamaissignaléeenAfrique, cf.Cremohipparionperiafricanum,deuxespècesderhinocéros:cf.Ceratotheriumsp.etaff.

Chilotheriumsp.,unProboscidiencf.Tetralophodonsp.,unGirafﬁdégen.etsp.indet.de grandetaillecomparableàcelledu«Palaeotragus»germainietdeuxbovidésdontl’unest probablementvoisindeProstrepsiceros,tandisquel’autreestunenouvelleantilopedetaille moyenneetàcornesspiralées,certainementunreprésentantdesCaprinae,groupetrèsrare enAfrique,oùaucuneformesemblablen’aétésignalée.C’estavecunâgeMiocènesupérieur correspondantplusprécisémentauTuroliend’Europe,quelafaunes’accordelemieux.

∗ Correspondingauthor.

E-mailaddress:[email protected](S.Zouhri).

doi:10.1016/j.crpv.2012.01.005

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456 S.Zouhrietal./C.R.Palevol11(2012)455–461 1. Introduction

As in other countries of North Africa, the Neogene macromammal faunas of Morocco are still very poorly documented.Infact,onlytwolocalitieshaveyieldedsig- niﬁcantremains.OneisthelateMiddleMiocenelocality ofBeniMellal(ChoubertandFaure-Muret,1961;Lavocat, 1961),theﬁrstimportantlocalityofMiocenevertebratesin Morocco,andtheotheristheLatePliocene/EarliestPleis- tocene localityof Ahl alOughlam, the mostproductive siteofthelateNeogeneofNorthAfricaandindeedoneof therichestoftheentireAfricancontinent(Geraads,2006;

Raynaletal.,1990).Mostothervertebratelocalitiesofthis periodhaveonlyyieldedmicromammals.Thediscoveryof largevertebrateremainsintheMioceneoftheOuarzazate basinis of greatpaleontologicaland geologicalinterest.

Indeed,whilethenewfossilshelptobridgeagapinthe NeogeneofNorthAfrica,andMoroccoinparticular,they alsoprovidesomeelementsofdatingthatwillhelptointer- pretthegeodynamicevolutionoftheregion.

2. Geographicalandgeologicalsetting

On the southern slope of the Central High Atlas in Morocco,Cenozoiccontinental depositsareknownfrom theOuarzazateforelandbasinandtheAïtKandoulaand AïtSedrat deposits (Fig. 1).Together, these continental deposits form the “Group of Imerhane” (Herbig, 1991) whichincludesfourcontinentalformationsranginginage fromtheLateEocenetotheQuaternary(ElHarﬁetal.,2001:

Fig.4).Theirgeologicalhistoryrepresentsamajorepisode ofsedimentationlastingfromtheEocenetotheQuater- nary,duringwhichalargebasinwasfragmentedandﬁlled withthickcontinentalseries,largelyduetotheupliftof thecentralHighAtlas(ElHarﬁetal.,1996,2001;Fraissinet etal.,1988;Gauthier,1960;GörlerandZucht,1986;Görler etal.,1988;Tesónetal.,2010).

TheOuarzazatebasinisanasymmetricsynclinelocated intheeasternextensionoftheSoussbasin,betweenthe mobileupliftareaoftheHighAtlasintheNorthandthesta- bledomainoftheAnti-AtlasintheSouth,ontheNorthern edgeoftheWestAfricanCraton.Itbelongstothestruc- turaldomainoftheAnti-Atlasandisseparatedfromthe HighAtlasbytheSouth-Atlasfault,whichistheeastern extensionoftheHayestransformfault.Thebasin,oriented onaneast-westaxis,is145kmlongandabout40kmwide atitsmaximuminthewesternpart(Fig.1).

Thisbasinisﬁlledwithcontinentalsedimentsranging fromtheUpperCretaceous(Senonian)totheQuaternary.

Paleogene deposits, mostly marine and lagoonal sedi- ments,areexposed almost exclusivelyin theNortheast oftheOuarzazate basin (Gauthier,1960).Herbig(1986, 1991) and Herbigand Trappe(1993) distinguishedﬁve formationsin this SubatlasicGroup. Their location was governedprincipallybytectonicbutalsobyeustaticcon- trols. In contrast,the Neogene deposits outcrop widely throughouttheOuarzazatebasin(Fig.1).Theycorrespond toamegasequenceoflacustrineandpalustrinesediments deposited in the context of an alluvial plain, and dot- tedwithlocalsabkhas.Theirthicknessvariesconsiderably fromthenorth,wheretheycanreach700m(Görleretal.,

1988)tothesouth,wherethesedimentsarebevelledonthe Anti-AtlasPrecambrianbasement.Theywereattributedto theAïtKandoulaFormation,originallydeﬁnedinthesmall basin ofthesamename(AïtKandoula,Fig.1)byGörler andZucht(1986)andGörleretal.(1988).Thisformation wasinitiallycomposedofthreemembersthathavebeen reducedtotwo(Fig.2)(ElHarﬁetal.,2001;seebelow).

Itislocallyexposedinverticaloutcropsintheimmediate vicinityoftheHighAtlaszone,anditbecomessubhorizon- talsouthwards.Thisformationliesunconformablyoverall olderstrataandinmanyplaces,forexampleonthesouth- ernedgesofthebasin,where thefossilsdescribedhere werefound;thedepositsﬁlltopographicdepressionscut intoolderstrata.Thedepositsofthisformationhavebeen subdividedintotwomembers(ElHarﬁetal.,2001):

•alowerpalustrineandlacustrinememberwithathick- nessofupto500m.Itconsistsoflimestone,travertine, marls,siltyclay,gypsumandsapropelite;

•an uppermember which consistsofalluvial deposits:

sandstoneandespeciallyconglomerates.

Thevertebratefossilsstudiedinthisworkarefromthe southernOuarzazatebasinbetweenthecityofSkouraand Imassinevillage,nearTiziN’Tadderht.

3. Paleontologicalbackground

IntheOuarzazatebasin,strataconsistinglargelyofPale- ogenedepositsinthenortheastofthebasinyieldedarich assemblageofmarineandcontinentalvertebrates,includ- inghundredsofspecies ofsharksand rays,bonyﬁshes, amphibians,turtles,crocodiles,lizards,snakesandmam- mals.Themammalsrepresentoneof theoldestrecords of the radiation of Eutheria in Africa. Gheerbrant et al.

(1992,1998)reviewedthesefaunasintheirstratigraphic and palaeogeographiccontext. Thelast majorpaleonto- logical discovery in the Ouarzazate basin was made in theLutetianlocalityofAznag(Tabuceetal.,2005)which yielded afaunaof mammalsandelasmobranchs associ- atedwithplanktonicforaminifera.UnlikethePaleogene, theNeogenedepositsthatoutcropwidelythroughoutthe Ouarzazate basin have only yielded ostracods, diatoms, algae, ﬁsh and micromammals, reported by Görler and Zucht(1986),Görleretal.(1988)andHelmdach(1988), whichsuggestaLanghiantoLowerSerravalianageforthe lowerpalustrineandlacustrineunitoftheNeogeneseries.

NorthoftheOuarzazatebasin,asetoffossilsitesdis- coveredintheAïtKandoulaBasinyieldcharophytes,ﬁshes, reptilesandmammals,andrangefromtheMiddleMiocene tothe Pliocene(Benammi,1997,2001; Benammiet al., 1995,1996;Görlerand Zucht,1986), whiletheremains of large mammals are rare and fragmentary. Benammi et al.(1995)and Rémyand Benammi(2006)reportthe presence, in various localities that have mostly yielded micromammals, of Gomphotheriidae indet., “Hipparion”

sp., Parapliohyraxsp., Sivatheriummaurusium, Girafﬁdae indet.,Gazellasp.,andaHippotragini,butthismaterialhas notbeendescribed.

Inthiswork,wereporttheﬁrstlargevertebrateassem- blagefromtheMiocene,discoveredintheregionofSkoura,

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Fig.1.GeologyandstructuralunitsofthesouthernslopesofCentralHighAtlaswesternpart(modiﬁedafterElHarﬁetal.,2001;Lavilleetal.,1977)and situationofthefossiliferouslocalities.

Fig.1. GéologieetunitésstructuralesdelapartieoccidentaleduflancsudduHautAtlascentral(d’aprèsElHarfietal.,2001;Lavilleetal.,1977modifié) etsituationdeslocalitésfossilifères.

inthesouthernpartoftheOuarzazatebasin.Itprovides aﬁrstinsightintothecompositionofthevariousfaunas, particularlythatoflargemammalsandtheirevolutionin theSouth-Atlasarea, andit alsocontributes tothedat- ingofsedimentaryand tectoniceventsthataffectedthe basin.TheprecisedatingofcontinentalFormationsofthe Ouarzazate basin iscritical tointerpreting thetectonic- sedimentaryevolutionofthearea,inparticularthelifting oftheHighAtlas.

4. Systematicpalaeontology

Thematerialwascollectedbyamateurcollectors,which naturally raises the problem of geographic and strati- graphicprovenance ofthefossils. Somenewspecimens have been collected by theauthors in December 2010.

These are the only ones whose origin is known with certainty (coordinates UTM zone 29, WGS84). Onepart of the material (about 20 specimens) is kept in a pri- vate museuminErfoud(Morocco),while theotherpart (about126specimens)ishousedattheFacultyofSciences AinChockofCasablanca(FSAC).Thefaunallistisasfol- lows:

Aves – cf. Struthio sp. shell fragments (FSAC) (739770/3450916).

Chelonia–cf.Centrochelyssp.Aperipheralplatefrom the right rear edge of the bridge; at the separation of

marginalscales,agroovereliefischaracteristicofterres- trialTestudinidae.Thesizeofthisplateisreminiscentof thegenusCentrochelys,whichisknownsincetheMiddle MioceneinnorthernAfricaandArabiabutisnowconﬁned totheSahel(LapparentdeBroin,2000).

Crocodylia – Crocodylus cf. niloticus Laurenti, 1768.

Analmost complete skull of a youngcrocodile (Fig.3I;

Erfoud), only lacking the basicranium and quadrate, is indistinguishablein itsmorphology and cranialpropor- tions,studiedindetail byTchernov(1986)andPickford (1996,2003),fromthemodernC.niloticus.Amongthefew noteworthydifferences,we notetheshortlengthofthe premaxillarelativetothewidthofthecranialtable(index 23/15Tchernov, 1986).Themedian nasalridge(“preor- bitalpromontorium”ofHecht,1987;“longitudinalnasal ridge”ofStorrs,2003),listedbyStorrsasacharacteristicof thisspeciesamongAfricancrocodiles,isbarelyvisiblehere, whichmaybeduetotheyoungageoftheanimal.Itappears thattheskullispartofthelineageoftheNilecrocodile,and mayevenrepresentthesamespecies.Thelatterappeared inAfricaintheLateMioceneinthelowermemberofthe NawataFormationatLothagam(Storrs,2003),andaccord- ingtoStorrs(2003),thecrocodilefromSahabi,C.checchia (Maccagno,1948;Hecht,1987)isalsoofthesamespecies.

Inthiscase,itispossiblethattheolderremainsfromJebel KrecheminTunisia,describedunderthatname(Geraads, 1989),shouldalsobereferredtoC.niloticus,pushingthe

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458 S.Zouhrietal./C.R.Palevol11(2012)455–461

Fig.2.SimpliﬁedlithostratigraphicsectionoftheImerhanecontinental GroupintheOuarzazatebasin.1:Mesozoiclimestoneandmarls(marine deposits);2:RedBeds(siltstoneandclaystonewithgypsumatthebase becomingsandstonesupward);3:CoarsesandstoneComplex(couplets ofsandstoneandmicroconglomeraticsfacies);4:Palustrineandlacus- trineComplex(palustrineandlacustrinelimestonewithevaporiticplaya deposits);5:Mio-Plioceneconglomerates(ﬂuvialandsiliciclasticalluvial fandeposits);6:Quaternaryconglomerates(conglomeraticpedimentand alluvialterracesdeposits).

Fig.2. Sectionlithostratigraphiquesimpliﬁée duGroupecontinental d’ImerhanedanslebassindeOuarzazate.1:calcairesetmarnesméso- zoïques(dépôtsmarins);2:couchesrouges(siltitesetargilitesàgypse àlabasedevenantgréseusesausommet);3:complexegréseuxgrossier (alternancedeniveauxgréseuxetmicroconglomératiques);4:complexe palustro-lacustre(calcairespalustro-lacustresetlocalementdépôtséva- poritiquesdeplaya);5:conglomératsmio-pliocènes(dépôtsﬂuviatiles etlocalementdecônesalluviaux);6:conglomératsquaternaires(dépôts deterrassesalluviales).

originofthespecies backtotheﬁrst partoftheUpper Miocene.

Mammalia – Equidae – Aff. Cremohipparion periafricanum (Villalta and Crusafont, 1957). A fragmentof mandible(Fig.3C;FSAC)is distinguishedfromall other specimensofHipparionsfoundatSkourabyitsverysmall size,reminiscentofthesmallSpanishspecies“Hipparion”

periafricanumplacedinthegenusCremohipparion(Zouhri

andBensalmia,2005).Thenewfragmentofmandiblehas a characteristicequinemorphologyand isclearlydiffer- entfromthose ofotherPerissodactylasuchastapirsor smallsuids.Itconsistsofthesymphysisandaportionof the leftanterior portionof the horizontalramus. How- ever,theincisorarchisdestroyed.Thehorizontalramus of themandibularis brokenontheleftside justbefore the front edgeof the secondpremolar (p2)and a little furtherforwardontherightside.Unfortunately,although some measures (7; 12;13 and 14 of Eisenmann et al., 1988:Fig.4A–B)ofthemandiblecanbeestimatedinthis specimen,itisimpossibletocompareitwiththeSpanish dwarfformswhosejawisunknown.Itisinanycasemuch smallerthantheotherknownsmallhipparionssuchasCre- mohipparionmatthewiAbel,1926; Cremohipparionnikosi BernorandTobien,1989or“Hipparion”elegansGromova, 1952.InNorthAfrica,theonlysmallhipparionsreported are“Hipparion”sitifense,nowconsideredanomendubium (BernorandScott,2003),andC.matthewiandC.nikosifrom Sahabi, recentlyidentiﬁed by Bernor et al. (2008)from postcranialmaterialintheﬁrstcase(C.matthewi),and a lower toothfragment inthe second.Irrespective ofthe taxonomicvalidityof“Hipparion”sitifense,materialasso- ciatedwiththistaxon(Eisenmann,1980)indicatesamuch morelargerspeciesthanthedwarfoneofTiziN’Tadderht.

TheTiziN’Tadderhthipparionshouldbeattributedtothe smallestknownhipparionsensulato,C.periafricanum,con- sidereduntilnowendemictoSpain,whereitisknownfrom numerousUpperTurolian(MN13)localities,asisthecaseof ValdecebroII(AlberdiandAlcala,1989-1990),Valdecebro 3,LasCasionesandElArquillo(VanDametal.,2001).

Cf.Hippotheriumprimigenium(VonMeyer,1829).The rest of the hipparion material from Tizi N’Tadderht is divided into two lots.The ﬁrst consistslargely of jugal teeth ofhipparions (mostof which belongto thesame individual),twoastragali,afragmentofMcIIIandsome phalanges(FSAC).Thesespecimensbelongtoalargeand heavilybuiltspecieswithaseriesofplesiomorphicchar- acters includingnotably brachyodont teeth,a lenticular protocone,andalackofthearticularfacetforthetrape- zoid onthe proximalarticularend of MC III. Thisform suggestsafﬁnitieswithHippotheriumKaup,1833,anOld WorldgroupknownthroughouttheUpperMiocene(Valle- sian andTurolian)andevenPliocene. Therobustnessof thepostcranialbonesandthedegreeofbrachyodontyof theuppercheekteethevokeH.primigenium(VonMeyer, 1829) rather than Hippotherium africanum Arambourg, 1959,anotablymoreslender-limbedandmorehypsodont species.

Hippotheriinigen.andsp.indet.Theotherpartofthe materialofTiziN’Tadderhthipparionsconsistofthreefrag- mentsofmandibles(Fig.3B),apartialMcIII,acompleteMT III,acalcaneumandafewphalanges(FSAC).Theybelong toanotablysmallerandmuchmoregracilespeciesthan thepreviousform(H.primigenium)withderivedcharac- ters suchas thepresence of a welldeveloped articular facet for thesmall cuneiformonthe proximalarticular surfaceofMtIII.Thetypicallyhipparioninedoubleknots and theabsenceofa developedectostylidonthelower cheek teethexcludetheassignment ofthis formtothe AfricangenusEurygnathohippusVanHoepen1930,which

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Fig.3. SomefossilsfromTiziN’Tadderht.A.Bovidaegen.etsp.indet.,skullinrightlateralview.B.Hippotheriinigen.andsp.indet.,rightlowertooth-row.

C.Aff.Cremohipparionperiafricanum,mandibularsymphysis.D.Aff.Chilotheriumsp.,P4.E–F.cf.Ceratotheriumsp.E.P4.F.Mandibleinleftlateralview.

G–H.Cf.Tetralophodonsp.G.Molarfragment.H.Mandible.I.Crocodyluscf.niloticus,skullinventralview.Scale=10cmforFigs.B–E;20cmforFigs.A,G,I;

40cmforFigs.F,H.

Fig.3. QuelquesfossilesduTiziN’Tadderht.A.Bovidaegen.etsp.indet.,crâneenvuelatéraledroite.B.Hippotheriinigen.andsp.indet.,rangéedentaire inférieuredroite.C.Aff.Cremohipparionperiafricanum,symphysemandibulaire.D.Aff.Chilotheriumsp.,P4.E–F.cf.Ceratotheriumsp.E.P4.F.Mandibuleen vuelatéralegauche.G–H.Cf.Tetralophodonsp.G.Fragmentdemolaire.H.Mandibule.I.Crocodyluscf.niloticus,crâneenvueventrale.Échelle=10cmpour lesFigs.B–E;20cmpourlesFigs.A,G,I;40cmpourlesFigs.F,H.

hasacaballinedoubleknotandalwaysanectostylidon thelowercheekteeth.Itisdifficulttodetermineeventhe supraspecific affinities of this material becausewe rec- ognize at present several genera of African hipparions, mainly defined by skull characters and the association betweenskull andpostcranialis notyet establishedfor mostofthesetaxa.Consequently,theHippotheriinimate- rial of Tizi N’Tadderht is attributed to a gen. and sp.

indet.

Rhinocerotidae–cf.Ceratotheriumsp.(FSC,Erfoud).This largerhinoceroswithrobustlegsisthemostcommonform and is probably linkedto the groupDicerotini because ofthedrasticreduction,ifnotabsence,ofincisorsinthe virtually complete mandible (Fig. 3F). This group, now exclusivelyAfrican,wascommonintheLateMiocenein theBalkano-Iranianprovince,whereitwasrepresentedby Ceratotheriumneumayri(orDicerosforsomeauthors).Itis alsodocumentedintheVallesianofAlgeriawith“Dicerorhi- nus”primaevusArambourg,1959(Geraads,1986),perhaps SahabiinLibya,butitsextensionfarthersouthinAfrica at this time is uncertain. A well-preserved MCIII from TiziN’Tadderhtis morerobustthanthatoftheAlgerian form, but lacks the strong concave side of C.neumayri (Giaourtsakis,2009),soaspeciﬁcdeterminationwouldbe toospeculative.

Aff.Chilotheriumsp.Asingleuppertooth(FSAC),cer- tainlyaP4,belongstoanotherrhinocerosspecies.Despite thewear,astronglypinchedprotolophalmostcompletely isolating the protocone, and a very long antecrochet interposed between the protocone and a reduced hypocone,distinguishitclearlyfromthatoftheDicerotini (compareFig.3DandE).Adetailedcomparisonisbeyond thescopeofthis note,but suchmorphologyisradically differentfromthatofDicerotini,butalsofromthatofthe brachypotheres,anothergroupknownintheLateMiocene inAfrica,andfromthatoftheelasmotheres,anotherset of Neogenerhinoceroses, where themedifossette tends tocloselingually.Thecharactersofthis tootharemore similartoaformthatGuérin(1966)haddescribedfrom theMioceneofTunisiaunderthenameDicerosdouariensis, andcompareevenbetterwiththeEurasianChilotherium, unknowninAfrica.Itgoeswithoutsaying,however,that suchdeterminationwouldbeinsufﬁcientlyfounded,and somewhatproblematicforbiogeographicalreasons.

Gomphotheriidae– Cf.Tetralophodonsp. Amandible (Fig. 3H;Erfoud)certainly had strongincisors but their exactsizeororientationcannotbedeterminedbecausethe symphysiswaslargelyreconstructedinplaster.Noteeth remain,withtheexceptionofasmallpartofthedentin oftherightm3.Thistoothhadtohave4or5thickand

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460 S.Zouhrietal./C.R.Palevol11(2012)455–461 transverselophids.Thelittlethatremains,however,shows

thatitwascertainlynotaformclosetotheUpperMiocene Stegotetrabelodon from Sahabi, whose lophids were far morenumerous,noraChoerolophodon,whoselophidsare arrangedinchevrons,noraformwithlophidsshowingan anancoidcontact.Thesecharactersareconﬁrmedbyafrag- mentoftooth(Fig.3G;FSC)withonehalfofarelativelythin lophidwithoutaccessoryconules.Themostlikelyhypoth- esisisthatthismaterialbelongstoaTetralophodon,ortoan Amebelodontidae,twotaxaknownintheMioceneofNorth Africa(Geraads,1989;Sandersetal.,2010).

Girafﬁdae– Girafﬁd gen.et sp.indet.Thisspecies is representedonlybyafewlimbbones(FSAC):adistorted distalend of a radius,a lunar (738283/3444137),and a cubo-navicular.Itis a formoflargesize,comparable to

“Palaeotragus”germainifromtheVallesianofBouHaniﬁa inAlgeria(Arambourg,1959),andprobablyalsopresentin BledDouarahandJebelKrechem(Geraads,1989).AllMid- dleMioceneGirafﬁdae,andparticularlythosefromNorth Africa(PalaeotraguslavocatiHeintz,1976fromBeniMellal) aremuchsmaller.

Bovidae–cf.Prostrepsicerossp.Thebaseofaslightly compressed bovid horn-core (FSAC) cannot belong to a gazelle because of its subtle but signiﬁcant twisting.

Some other fragments correspond in size to the same form,whichshouldprobablybereferredtoataxonclose toProstrepsiceros,a genusknownmainlyin theeastern Mediterranean,butalsoreportedfromSahabi(Lehmann andThomas,1987)andJebelKrechem(Geraads,1989).It isnotcertain,however,thattheNorthAfricanformsreally belongtothesamegenusastheonesfromtheBalkano- IranianProvince(BouvrainandBonis,2007).

Bovidaegen. etsp. indet.Askull of a medium-sized antelope(Fig.3A;Erfoud),completewiththeexceptionof theanterioranddorsalpartsoftheface,isprobablythe mostinterestingfossilfromTiziN’Tadderht.Theskullhas longhornsthat areslender,divergent,fairlytightlyspi- raledwithahomonymoustorsion,therebyexcludingany relationshipwithmostspiralhornedantelopesfromthe Miocene.Therearoftheskullisveryinclinedwithrespect tothefacialregion,thetuberositiesofthebasioccipitalare broadandﬂattened,andthedentitionis“aegodont”.This showsthatitiscertainlyarepresentativeoftheCaprinae.

ThisgroupiscommonintheLateMioceneBalkano-Iranian province,butthehornsareneverspiraled,andveryrarein Africa,wherenosimilarformhasbeenreported.

5. Conclusions:ageoffauna

Thepreciseoriginofmostofthefossilsidentiﬁedabove isunknown;theircontemporaneityisnaturallyinques- tion. However, there is no evidence that the fauna is heterochronous,althoughtheextentandthicknessofthe uppermemberoftheAïtKandoulaFormationmakeitdif- ﬁcultto establish the strict contemporaneityof all the elements.Still,there isnomajorchangein depositional environmentswithinthismember,anditisunlikelythat itdocumentsaverylongperiodoftime,assuchdetritic depositsmayaccumulatequiterapidly.Paradoxically,itis a“reptile”thatcanhardlybedistinguishedfromanextant form,whichmightprovidethestrongestargumentinfavor

ofa heterochronyoftheassembly,buttheagerangeof C.niloticusmightstretchbacktoatleastthesecondhalfof theMiocene(TurolianmammalageinEurope).Thisageis themostconsistentwiththerestofthefauna.Thepres- enceofahipparionnaturallyexcludesanageolderthan theUpperMiocene,whilethelargesizeofthegirafﬁd,the massivenessofthemainbonesoftherhinoceros,andthe derivedfeaturesofthenewbovidareconsistentwithfau- nasofthisage,ratherthanwitholderfaunas(Vallesian).

Themainargumentthatwouldsuggestapost-Mioceneage isthetotalabsenceofanthracotheres.Indeed,thisgroup ispresentinmostdepositsoftheupperMioceneinCen- tralAfrica(Toros-MenallainChad),Tunisia(BledDouarah, JebelKrechem)andLibya(Sahabi).However,itisabsent in BouHaniﬁa inAlgeria and in themiddleMiocene of BeniMellalinMorocco,andwecanthereforehypothesize thatitwasrelatedtoawatersystemthatdidnotextend tothewest (Lihoreau etal.,2006),and itsabsencein a Moroccansitewouldbeofnochronologicalsigniﬁcance.

Thescarcityof“Turolian”depositsinNorthAfricaisobvi- ouslyanobstacletotheprecisedatingoffaunasoftheAït KandoulaFormation.

Althoughitisrepresentedbyasmallsample,thefauna ofTiziN’Tadderhtisinterestinginseveralaspects.Besides itscontributiontothedatingoftheuppermemberofthe AïtKandoulaFormation,itlinksolderMiocenefaunasfrom Africa (Beni Mellal and Vallesian faunas of Algeria and Tunisia)withmorerecentones(Sahabi).Thepresenceofan unusualantelopehintsathowmuchremainstobelearned aboutmammalianfaunasfromthelateNeogeneofAfrica.

Acknowledgments

The authorsthank MrBrahim Tahiri for allowingus tostudysomeofthematerialsintheMuseumofErfoud, GeorgiN.Markov,andFrancedeLapparentfortheirhelpon theProboscideaandthetortoiseofTiziN’Tadderht,respec- tively.Wealsothanktwoanonymousreviewersfortheir usefulcomments.

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