an early cretaceous vertebrate assemblage from the cabao ... · strongly differs from both the...

Geol Mag 147 (5 ) 2010 pp 750ndash759 ccopy Cambridge University Press 2010 750doi101017S0016756810000178

An Early Cretaceous vertebrate assemblage from theCabao Formation of NW Libya

JEAN LE LOEUFFlowast EDDY METAISdagger DIDIER B DUTHEILDagger JEAN LOUP RUBINOsectERIC BUFFETAUTpara FRANCOIS LAFONTsect LIONEL CAVIN FABRICE MOREAU

HAIYAN TONGlowastlowast CHRISTIAN BLANPIEDdagger amp ALI SBETAdaggerdaggerlowastMusee des dinosaures 11260 Esperaza FrancedaggerTotal Tour Coupole Paris la Defense France

Dagger15 passage du Buisson Saint-Louis 75010 Paris FrancesectTotal CSTJF Avenue Larribau 64018 Pau Cedex France

paraCNRS Ecole Normale Superieure 24 rue Lhomond 75231 Paris cedex 05 FranceMuseum drsquoHistoire Naturelle 1 route de Malagnou 1208 Geneve Switzerland

Domaine de la Grange 37110 Les Hermites Francelowastlowast30 rue Carnot 94270 Le Kremlin-Bicetre FrancedaggerdaggerGeology Department University of El Fateh Tripoli Libya

(Received 21 November 2009 accepted 18 January 2010 First published online 10 March 2010)

Abstract ndash Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwesternLibya include the hybodont shark Priohybodus the crocodilian Sarcosuchus an abelisaurid abaryonichine spinosaurid and a large sauropod with spatulate teeth The Cabao Formation may beHauterivian to Barremian in age although an earlier Berriasian to Valanginian age cannot be excludedIts dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger andstrongly differs from both the Cenomanian assemblages of Morocco and Egypt and the Late Aptianto Albian fauna of Tunisia Fossil vertebrates may be an important tool to establish the stratigraphicalframework of the poorly dated Early Cretaceous continental deposits of Africa

Keywords Early Cretaceous Libya dinosaurs biostratigraphy

1 Introduction

Early Cretaceous vertebrates are still poorly known inAfrica where the most important Cretaceous localitiesare Cenomanian in age in Egypt and Morocco (Stromer1936 Smith et al 2001 Cavin et al 2010 Taquet1976) Earlier sites of the lsquoContinental Intercalairersquo (aname suggested by Kilian in 1931 to designate thePalaeozoic and Mesozoic continental levels locatedbetween the latest marine Palaeozoic rocks and theCenomanian transgression across the Sahara desert)in Niger (Taquet 1976 Sereno et al 1994) Mali(OrsquoLeary et al 2004) Algeria and several other places(see Rauhut amp Lopez-Arbello 2009 for a recent review)are still extremely poorly dated because of the absenceof marine incursions during the early Cretaceous inthese areas Other Early Cretaceous sites have beenrecognized in Tunisia (Bouaziz et al 1988 Bentonet al 2000 Srarfi et al 2004 D Srarfi unpub PhDthesis Univ Claude Bernard Lyon 2006) where somemarine intercalations can help to pin down their age

In the course of sedimentological field work forTOTAL Libye one of us (EM) discovered severalrich assemblages of fossil vertebrates in the CabaoFormation in NW Libya (surface collecting) Thismaterial is described herein The discovery of ante-Albian vertebrates in Libya is of great interest for our

lowastAuthor for correspondence jeanleloeuffyahoofr

understanding of the succession of early Cretaceousvertebrate assemblages in Africa as some correlationscan be made with the neighbouring assemblages ofSouthern Tunisia the age of which is discussed below

2 A short history of true and mythical discoveries ofvertebrates in the Early Cretaceous of Libya

According to several authors (Robaux et al 1942Lapparent 1951 1960 Ouaja et al 2002 Srarfi et al2004 D Srarfi unpub PhD thesis Univ ClaudeBernard Lyon 2006) Early Cretaceous continentalvertebrates were first recognized in Libya by Italiangeologists who described fossil fishes in the depositsof the lsquoContinental Intercalairersquo This mention oflsquoItalian authorsrsquo can be traced originally to Robauxet al (1942) and was then cited by later authorsIn this paper (the notice of a hydrogeological mapof southern Tunisia original pamphlet undated andperhaps effectively published in 1942 or 1943) Robauxet al mention the recent discovery of a lsquorich fish faunamade in Tripolitania and recently described by Italianauthors which leaves no doubt about the Wealden age ofthese deposits these fishes are as followsrsquo (translationfrom the French by the authors) but the text stops herewithout any list following the colon In a long erratumkept at the Societe geologique de France in Paris it isnoted that this publication was made difficult by the

httpswwwcambridgeorgcoreterms httpsdoiorg101017S0016756810000178Downloaded from httpswwwcambridgeorgcore University of Basel Library on 11 Jul 2017 at 071646 subject to the Cambridge Core terms of use available at

Early Cretaceous vertebrates from Libya 751

lsquo1943 Tunisian eventsrsquo which prevented the authorsfrom correcting the proofs The so-called events arerelated to some episodes of the Second World Warin North Africa when German and Italian troopsinvaded the French protectorate of Tunisia at the endof 1942 before they had to leave the country in 1943following the battle of El Alamein Their defeat led tothe departure of Italians from Libya and the closureof their geological institutions in Africa Howeverwe could not find a single original reference in theAnnali del Museo Libico di Storia Naturale about thesepurported discoveries of fishes and Desio (1950) in hisbibliography of the geology of Libya does not mentionEarly Cretaceous (lsquoWealdenrsquo) fossil fishes found byItalian geologists It thus seems likely that Robaux et alwould have heard about discoveries of fossil fishes inTripolitania by Italian colleagues which were neverpublished and soon forgotten following the collapseof Italian colonies in North Africa An alternativepossibility is that Robaux et al mistakenly mentionedfishes when referring to Sassirsquos recent (1942) reportof Wealden plants in the Early Cretaceous of LibyaAlthough a forgotten publication in some obscurejournal cannot be excluded we suspect that one ofour hypotheses is correct Lapparent was thus the firstauthor to report Early Cretaceous vertebrates fromLibya in 1960 he mentioned several sites discoveredby him along the Dahar escarpment in South Tunisiaand Tripolitania (northwestern Libya) during a fieldtrip in 1951 on the tracks of the French geologistPervinquieres who was the first in 1912 to describefossil vertebrates from the same horizons in southernTunisia In Libya Lapparent mentions several localitiesin Nalut Giado Iefren and Garian The Giado localityyielded a theropod tooth and a bone referred by himto Elaphrosaurus iguidiensis Other sites providedfishes and crocodiles only and were not described indetail by Lapparent Later Burollet amp Manderscheid(1965) mentioned teeth and jaws of crocodiles fromthe Cabao Formation Hammuda mentioned in 1969 acrocodile tooth found at Wadi al-Mezaida near GiadoThe Libyan geologist El-Zouki (1980) later figuredvarious teeth and scales referred to fishes turtles andcrocodiles as well as a dinosaur vertebra from the CabaoFormation in a sand quarry at Jannawan near GiadoAlthough El-Zouki refers this material to Spinosaurus() the published plate (a vertebral centrum in ventralview) does not allow any precise assessment Morerecently Smith amp Dalla Vecchia (2006) describeda possible abelisaurid tooth collected by an Italianamateur palaeontologist near Nalut (see also DallaVecchia 1995 for a preliminary description of thistooth) The same locality has yielded abundant materialof the hybodont shark Priohybodus described by Duffinin 2001 This material has been found according toDuffin (2001) and Smith amp Dalla Vecchia (2006) in theKiklah Formation which overlies the Cabao Formationbut our own investigations in this area suggest thatthis material comes rather from the Cabao FormationSmith et al (2006) and Smith amp Lamanna (2009) in

Figure 1 Location map

two abstracts briefly reported more discoveries in theCabao and Kiklah formations including abelisauroidbones a possible titanosauriform along with sharkturtle lungfish and crocodyliform remains

3 Geology

Several sand quarries located along the northernborder of the Jabal Nafusah have yielded vertebrateremains The material described here was found in threequarries Tendamirah Jawsh and Jannawan (Fig 1)These are all opened in the Cabao Formation acontinental to shallow marine unit (estuarine with tidalinfluences on depositional settings cf Rubino et al2009) first described by Burollet in 1963 (Burollet1963ab) near the village of Giado At this localityit directly overlies the marine Late Jurassic ShakshukFormation and underlies the Kiklah Formation (Fig 2)a possible lateral equivalent of the Oum ed Diab andChenini members of the Aın Guettar Formation ofsouthern Tunisia considered as Late Aptian to Albianin age (see Ouaja et al 2002) Since that time thisstratigraphic succession has been adopted by all authorsworking in Djebal Nefusah (Hammuda 1969 El-Zouki 1980 Fatmi Eliagoubi amp Hammuda 1980)but eight sedimentological sections logged betweenthe Giado locality and the Tunisian border clearlydemonstrate that an intermediate sandy and shalyinterval appears and develops between the Shakshukand Cabao formations in this area (Fig 3) This seriesup to 120 m thick is topped by a distinct memberof lagoonal green shales and dolomites that clearlycorresponds to the Douiret shale first described insouthern Tunisia by Busson in 1967 and ranged morerecently in the upper Douiret Formation by Ouajaet al (2002) This additional interval intercalatedbetween the Shakshuk and the Cabao formations isherein mentioned for the first time in Libya andshould be considered as the lateral equivalent ofthe Tunisian Boulouha and Douiret formations (see


752 J LE LOEUFF AND OTHERS

Figure 2 Composite stratigraphical section of the Jabal Nafusah NW Libya



Figure 3 Lithostratigraphical correlation between Jannawan Wazzin (TunisianndashLibyan border) and Dehibat (SE Tunisia) sectionsDehibat section is from Ouaja et al 2002 (modified) Horizontal aspect not to scale Distance between Dehibat and Jannawan 120 km

Dehibat section modified from Ouaja et al 2002 inFig 3) The overlying Cabao Formation would itselfbe contemporaneous with the sandstones overlying theDouiret shales in Tunisia and appearing south of Touilel Hira (lsquoBarremian sandstonesrsquo of Busson 1967) andlater attributed to the Chenini Formation by Bouazizet al (1988) and Ouaja et al (2002) We suspect thatonly the uppermost part of this sandy interval shouldbe considered as the AptianndashAlbian Chenini Formation(see Fig 3)

Tunisian geologists refer the Douiret Formation tothe Aptian or Barremo-Aptian This age assignment isbased on its geometric position above Hauterivian toBarremo-Bedoulian levels of the Boulouha Formation(the latter being dated by the occurrence of the LateHauterivian to Early Aptian (Bedoulian) rhynchonellid

Loriolithyris russillensis cf Peybernes et al 1996) andbelow the Chenini member of the Upper Aptian toAlbian Aın Guettar Formation The Douiret Formationis thus generally considered as Barremo-Aptian in ageby Tunisian geologists (Ouaja et al 2002 Srarfi et al2004) However the brachiopod L russilensis has alsobeen described from older Berriasian strata in Algeria(Benest Gaspard amp Gali 1996) which would rathersuggest an age somewhere between the Berriasian andthe Early Aptian for the Boulouha and the Douiretformations and the overlying lowermost sandstoneinterval in Tunisia equivalent of the Cabao FormationThe fossil content of the Douiret Formation can helpto constrain its age the absence of angiosperms inits flora (Barale amp Ouaja 2002) would also suggestan age older than Aptian The microvertebrates from



the Douiret Formation are known from the LateJurassic to the Aptian the youngest record of severaltaxa being in the Hauterivian or Barremian (GCuny pers comm) the latest ascertained record ofEgertonodus is in the Barremian (Rees amp Underwood2008) Caturus has been found in the Barremian ofSpain (Buscalioni et al 2008) but not later and theyoungest record of Gyrodus is in the Hauterivianof Germany (Kriwet amp Schmitz 2005) This faunalassemblage thus strongly suggests a pre-Aptian age forthe Douiret Formation the macrofauna does not yielduseful stratigraphical information In all likelihooda Hauterivian to Barremian age is plausible for theDouiret Formation although an older Berriasian orValanginian age cannot be completely excluded

4 Vertebrates from the Cabao Formation

The material described in this paper belongs tothe collections of the Geology Department at theUniversity Al Fateh in Tripoli Libya (abbreviationUT)

4a Sharks and bony fishes

The fish assemblage is composed of hybodonts (Hy-bodus sp Priohybodus arambourgi) actinopterygians(Semionotiformes indet) and sarcopterygians withone dipnoan (Ceratodontidae a large tooth) and amawsoniid coelacanth Teeth of the hybodont sharkPriohybodus arambourgi are extremely abundant inmost localities of the Cabao Formation whereas thisspecies is conspicuously absent from the overlyingKiklah Formation Priohybodus is known only fromEast and Northeast Africa (Somalia Ethiopia TunisiaLibya) in the Late Jurassic and the Early Cretaceousbut is unknown from the contemporaneous localitiesof West Africa This might suggest some kind ofendemism for this freshwater hybodont but its recentdiscovery in probably Late Jurassic levels in Uruguay(Perea et al 2001) rather suggests that its absence inNiger or Morocco may reflect stratigraphical andorenvironmental differences P arambourgi might be re-stricted to a Late Jurassicndashearliest Cretaceous interval

The tooth plate is referable to Ceratodus cf humeibecause it shows few crests (probably five) and thecrests are very shallow This species which may reach avery large size is known in the Cretaceous lsquoContinentalIntercalairersquo of southern Morocco eastern and southernAlgeria Niger Mali Sudan and Egypt (Tabaste 1963Martin 1984 Werner 1994 Churcher amp De Iuliis2001 Churcher De Iullis amp Kleindienst 2006)

Some ganoıd scales and circular teeth could bereferable to some Semionotiformes fishes This actino-pterygian group is widely distributed in the Cretaceouscontinental beds of Africa

Several fragments showing a strong reticulatedornamentation can be referred with confidence topieces of the skull roof and opercular series of amawsoniid coelacanth A fragment (JAN-7) of a bony

plate bearing irregularly arranged small bulbous teeth isreminiscent of the toothed parasphenoid of Mawsonialavocati from the Kem Kem beds in Morocco figuredby Wenz (1981) and Cavin amp Forey (2004) or of thetoothed pterygoid of Mawsonia gigas from the EarlyCretaceous of Brazil figured by De Carvalho amp Maisey(2008)

4b Turtles and crocodiles

El-Zouki (1980) referred a xiphoplastron from Jan-nawan Quarry to an undetermined pleurodiran Manyisolated crocodilian teeth cannot be referred to knowngenera however several morphotypes can be distin-guished Some very large teeth from the Tendamirahquarry have a smooth enamel and may belong to thegiant pholidosaurid Sarcosuchus imperator from theEl Rhaz Formation of Niger (Broin amp Taquet 1966Taquet 1976 Sereno et al 2001) Sarcosuchus is alsoknown from Algeria and from the Douiret Formationof Tunisia as well as from the Early Cretaceous(not later than Aptian) of Brazil (Buffetaut amp Taquet1977)

4c Dinosaurs

Only Saurischian dinosaurs have been found in theCabao Formation No ornithopod teeth have been foundso far during our field work although it would bepremature to conclude that they are absent in theCabao Formation given the limited size of our sampling(surface collecting) It should be noted that one isolatediguanodont tooth has been mentioned in the DouiretFormation of Tunisia by Srarfi et al (2004) Smith ampDalla Vecchia (2006) referred an isolated tooth fromNalut to the family Abelisauridae on the basis of amorphometrical analysis

Two teeth from the Jawsh quarry share severalvery peculiar characters The best preserved specimen(UT-JAW2 Fig 4) is a 27 mm high recurved toothslightly compressed labio-lingually with a granularenamel The tooth is fluted and its mesial and distalcarinae show extremely small serrations (7ndash8mm)Their morphology is very close to that of the teeth of thespinosaurid Baryonyx from the Barremian of England(Charig amp Milner 1997) although they seem to bebroader in relation to the crown height than is typical forBaryonyx We refer them to the sub-family Baryonych-inae Baryonychine spinosaurids have been describedfrom the El Rhaz Formation in Niger Suchomimustenerensis Sereno et al 1998 being probably a juniorsynonym of the extremely incomplete Cristatusauruslapparenti Taquet amp Russell 1998 according toMilner (2003) however the spinosaur from Niger isindistinguishable from Baryonyx walkeri Although theauthors of these two papers did not fully illustrate theteeth of these dinosaurs they mention that they bearextremely small serrations The presence of serrationson the teeth is a good diagnostic character to separatebaryonychine spinosaurids which have such serrations



Figure 4 UT-JAW2 tooth of cf Baryonyx (Jawsh Quarry Cabao Formation Libya) in (a) labial (b) distal (c) lingual and (d) mesialviews (e) Enlargement of the mesial carina Scale bars 10 mm (andashd) and 2 mm (e)

Figure 5 UT-TEN15 tooth of a camarasaurid sauropod (Tendamirah Quarry Cabao Formation Libya) in (a) mesial (b) labial (c)lingual and (d) distal views scale bar 20 mm

from spinosaurine spinosaurids (that is Spinosaurusaegyptiacus Irritator challengeri) the teeth of whichhave no serrations In Africa Spinosaurinae occurin the youngest formations of the lsquoContinental Inter-calairersquo that is in Albian and Cenomanian depositsfrom Egypt Algeria Tunisia and Morocco whereasolder ante-Albian spinosaurids which are known fromNiger and Cameroon (J D Congleton unpub Masterrsquosthesis Southern Methodist Univ 1990) all belong tothe Baryonichinae suggesting a replacement duringthe Late Early Cretaceous (Buffetaut amp Ouaja 2002)Buffetaut (2008) has mentioned possible spinosauridteeth with larger serrations from the Late Jurassic ofTanzania

A single tooth from the Tendamirah Quarry (UT-TEN15 Fig 5) can be referred to a sauropod dinosaurIt is a large (preserved height 74 mm) spatulate toothwith a wrinkled enamel the crown of which is brokenat mid-height There is a distinct neck between the

root and the crown The labial face is convex andthe lingual face is flat with two coalescing centralridges delimiting a mesial and a rostral concavityThe rostral and mesial edges of the tooth are paralleland vertical This tooth (which belongs to the spoon-like tooth morphotype of Calvo 1994) is clearlydifferent from the teeth of Brachiosauridae whichshow no constriction between the crown and the rootand is also different from the various morphotypesillustrated by Srarfi (D Srarfi unpub PhD thesisUniv Claude Bernard Lyon 2006) from the Cheninimember in Tunisia This constriction is reminiscentof the Turiasauria (cf Royo-Torres Cobos amp Alcala2006) but the crown does not show the typical heart-shape of this poorly known clade This tooth hasthe closest affinities with those of Camarasaurusfrom the Late Jurassic of the United States aswell as with teeth from the Tiouraren Formation ofNiger initially described by Lapparent (1960) under



the name of Rebbachisaurus tamesnensis and laterreferred to Jobaria tiguidensis by Sereno et al (1999)although the figured teeth of Jobaria are markedlydifferent (see next Section) We provisionally refer thetooth from Tendamirah to the family Camarasauridae(Camarasauridae indet) Although Rich Molnar ampRich (1983) mentioned camarasaurid teeth from theLate JurassicndashEarly Cretaceous Kirkwood Formation inSouth Africa their figures do not show any constrictionat the base of the crown and their material rathersuggests a brachiosaurid sauropod

5 Biostratigraphical implications of the vertebrateassemblage

The ubiquitous abundance of the hybodont Pri-ohybodus arambourgi may be interesting for biostrati-graphical purposes as the genus is only known fromLate Jurassic levels outside Libya and Tunisia (that is inYemen Uruguay Somalia and Ethiopia) Its last occur-rence indeed was supposed to be in the overlying KiklahFormation (see Duffin 2001) but our field work at thesite which has yielded the material studied by Duffinas mentioned above shows that it was actually foundin the Cabao Formation not in the Kiklah FormationIt is extremely abundant in the Cabao Formation butconspicuously absent in the Kiklah Formation whichis believed to be of Albian age (Novovic 1977 Tekbali2000) It can be noted that this genus is also the mostfrequent hybodont in the Tunisian Douiret Formationwhereas a much more diversified shark fauna is knownfrom the Chenini and Oum ed Diab members of LateAptian to Albian age (Cuny et al 2004) even thoughthis may be linked to environmental differences asPriohybodus is considered a freshwater form it isinteresting to note that it is absent from the Tunisianformations of supposed AptianndashAlbian age and that itslast well-dated occurrence is in the Douiret Formation(Cuny et al 2004) that is in the Berriasian to EarlyAptian of Tunisia The record of Priohybodus wouldthus instead suggest an age close to the beginning ofthe Cretaceous for the Douiret and Cabao formations

The presence of a baryonichine spinosaurid alsosuggests an ante-Albian age as African baryonichinesare so far known only from the supposedly Aptian ElRhaz Formation of Niger and the Koum Formation ofCameroon the vertebrate assemblage of which seemsvery close to the El Rhaz fauna (Taquet 1984 J DCongleton unpub Masterrsquos thesis Southern MethodistUniv 1990) European baryonichines are known fromthe Hauterivian to the Early Aptian in England Portugaland Spain (Milner 2003 Buffetaut 2007 Canudo et al2008) The rich Albian and Cenomanian deposits ofNorth Africa have so far yielded only spinosaurinespinosaurids (Cavin et al 2010 Buffetaut amp Ouaja2002) and spinosaurids are so far unknown from theEuropean Albian localities The precise age of theEl Rhaz Formation is not so clear it was consideredas Barremian to Late Albian by Taquet (1976) whosuggested an Aptian age an opinion largely followed

by subsequent authors although Taquetrsquos suggestionwas based on the referral to the Aptian by Busson(1967) of an isolated ornithopod tooth found in Tunisiaby Lapparent It has since been demonstrated that thisiguanodontid tooth (found in the Oum ed Diab memberof the Ain Guettar Formation a few metres below theCenomanian marine deposits) was more likely Albianafter the discovery of ammonites in the vicinity (DSrarfi unpub PhD thesis Univ Claude Bernard Lyon2006) However iguanodontid teeth are also knownin the older Douiret Formation (Srarfi et al 2004)Thus Taquetrsquos main argument for an Aptian age seemsweaker and his lsquoBarremian to Albianrsquo interval for theEl Rhaz Formation is likely although an older age isnot impossible as the lower part of the formation is notstratigraphically constrained

The presence of a sauropod spatulate tooth in Libyawould also suggest an age older than Aptian for theCabao Formation Although the best known camara-saurid genera (Camarasaurus and Haplocanthosaurus)are restricted to the Late Jurassic of the UnitedStates isolated bones and teeth from Europe areslightly younger (Berriasian of Western France cfLe Loeuff Buffetaut amp Merser 1996 Colin et al2004) The youngest possible camarasaurids knownso far are Hauterivian to Early Barremian in age inEurope (locality of El Pelejon in Galve Spain cfSanz et al 1987) incomplete spatulate teeth occurin the Tiouraren Formation of Niger which underliesthe El Rhaz Formation and the age of which hasbeen strongly disputed by Rauhut amp Lopez-Arbarello(2009) who on the basis of the evolutionary grade ofthe dinosaurs suggest a Late Middle Jurassic age forthis unit El Rhaz sauropods include the rebbachisauridNigersaurus taqueti and an undescribed titanosaur(Taquet 1976 Sereno et al 1999) but spoon-liketeeth have been reported only in the older TiourarenFormation (see Lapparent 1960) Sereno et al (1999)have described the sauropod Jobaria in this formationthe single illustrated tooth shows serrations and ismarkedly different from the Libyan tooth and alsofrom the spoon-like teeth illustrated by Lapparent thissuggests that the spoon-like teeth from Niger may notbelong to Jobaria but to a second sauropod possiblya camarasaurid The absence in the Cabao Formationof ornithopod teeth which are extremely abundant inthe El Rhaz Formation (but absent in the TiourarenFormation) can hardly be taken into consideration giventhe small size of our sampling

6 Conclusions

The vertebrate assemblage from the Cabao Formationthus strongly suggests an Ante-Aptian age possiblyHauterivian to Barremian like the Douiret Formation ofTunisia It can be suggested that the El Rhaz Formationof Niger which shows some vertebrates in commonwith the Cabao Formation may be slightly youngerthat the latter (absence of Priohybodus and sauropodswith large spatulate teeth abundance of iguanodontids)



and that the Tiouraren Formation may be slightly olderbut possibly still Cretaceous (presence of sauropodswith spatulate teeth) In Tunisia the younger AinGuettar Formation (AptianndashAlbian) has yielded avery different dinosaur assemblage (spinosaurinesadvanced titanosauriforms) This review confirms thestrong potential of fossil vertebrates to correlatesites in the Continental Intercalaire across AfricaContrary to what was initially suggested by Lappar-ent (1960) who envisioned a rather uniform faunathroughout these deposits it is now widely recognizedthat important changes occurred in the vertebrateassemblages between the earliest Cretaceous and theCenomanian Although not enough assemblages havebeen described in detail it seems that vertebrates maybe useful to establish the stratigraphical frameworkof the Continental Intercalaire and its equivalentsin central eastern and northern African Cretaceousbasins

Acknowledgements We wish to thank the NOC of Libya forhaving authorized TOTAL to perform the sedimentologicalfield work and NOC and TOTAL Libye for permissionto publish this paper We thank Gilles Cuny (GeologicalMuseum Copenhagen) for sharing information on the ageof the Tunisian Douiret Formation The senior author thankscolleagues from the Office National des Mines in Tunis (MBel Hadj M Ouaja D Srarfi) for their invitations and jointfield work in southern Tunisia Thanks to Didier Cave for thedrafting of the Geology section and to Gilles Cuny and ananonymous reviewer for their constructive remarks

References

BARALE G amp OUAJA M 2002 La biodiversite vegetale desgisements drsquoage Jurassique superieur-Cretace inferieurde Merbah el Asfer (Sud-Tunisien) Cretaceous Re-search 23 707ndash37

BENEST M GASPARD D amp GHALI M 1996 Lesbrachiopodes de lrsquoavant-pays ouest-algerien lors dela phase transgressive maximale du Berriasien super-ieur systematique environnement et paleogeographieGeobios 29 13ndash34

BENTON M J BOUAZIZ S BUFFETAUT E MARTILL DOUAJA M SOUSSI M amp TRUEMAN C 2000 Dinosaursand other fossil vertebrates from fluvial deposits inthe Lower Cretaceous of southern Tunisia Palaeogeo-graphy Palaeoclimatology Palaeoecology 157 227ndash46

BOUAZIZ S BUFFETAUT E GHANMI M JAEGER J-JMARTIN M MAZIN J-M amp TONG H 1988 Nouvellesdecouvertes de vertebres fossiles dans lrsquoAlbien du SudTunisien Bulletin de la Societe geologique de France 4335ndash9

DE BROIN F amp TAQUET P 1966 Decouverte drsquouncrocodilien nouveau dans le Cretace inferieur du SaharaComptes Rendus de lrsquoAcademie des Sciences de Paris D262 2326ndash9

BUFFETAUT E 2007 The spinosaurid dinosaur Baryonyx(Saurischia Theropoda) in the Early Cretaceous ofPortugal Geological Magazine 144 1021ndash5

BUFFETAUT E 2008 Spinosaurid teeth from the LateJurassic of Tendaguru Tanzania with remarks on theevolutionary history and biogeographical history of theSpinosauridae In Mid-Mesozoic Life and EnvironmentsCognac (France) (eds J-M Mazin J Pouech P

Hantzpergue amp V Lacombe) pp 26ndash8 Documents desLaboratoires de Geologie de Lyon 164

BUFFETAUT E amp OUAJA M 2002 A new specimen ofSpinosaurus (Dinosauria Theropoda) from the LowerCretaceous of Tunisia with remarks on the evolutionaryhistory of the Spinosauridae Bulletin de la Societegeologique de France 173 415ndash21

BUFFETAUT E amp TAQUET P 1977 The giant crocodilianSarcosuchus in the Early Cretaceous of Brazil and NigerPalaeontology 28 203ndash8

BUROLLET P F 1963a Saharan symposium Field trip Guidebook of the excursion to Jebel Nefusa Tripoli PetroleumExploration Society of Libya 17 pp

BUROLLET P F 1963b Discussion sur la StratigraphieLibyenne Revue de lrsquoInstitut Francais du Petrole Paris18 1323ndash1328

BUROLLET P F amp MANDERSCHEID G 1965 Le Cretaceinferieur en Tunisie et en Libye Memoires BRGM 34785ndash94

BUSCALIONI A D FREGENAL M A BRAVO A POYATO-ARIZA F J SANCHIZ B BAEZ A M CAMBRA

MOO O MARTIN CLOSAS C EVANS S E ampMARUGAN LOBON J 2008 The vertebrate assemblageof Buenache de la Sierra (Upper Barremian of Serraniade Cuenca Spain) with insights into its taphonomy andpalaeoecology Cretaceous Research 29 687ndash710

BUSSON G 1967 Le Mesozoıque saharien 1ere partielrsquoextreme Sud-Tunisien lsquoCentre Recherche Zones ar-idesrsquo Centre National de la Recherche Scientifique editParis 8 194 pp

CALVO J O 1994 Jaw mechanics in sauropod dinosaursGaia 10 183ndash93

CANUDO J I GASULLA J M GOMEZ-FERNANDEZ DORTEGA F SANZ J L amp YAGUE P 2008 Primeraevidencia de dientes aislados atribuidos a Spinosauridae(Theropoda) en el Aptiano inferior (Cretacico Inferior)de Europa Formacion Arcillas de Morella (Espana)Ameghiniana 45 649ndash52

CAVIN L amp FOREY P L 2004 New mawsoniid coelacanth(Sarcopterygii Actinistia) remains from the Cretaceousof the Kem Kem beds SE Morocco In Mesozoic FishesIII ndash Systematics Paleoenvironments and biodiversity(eds A Tintori amp G Arratia) pp 493ndash506 Dr PfeilVerlag

CAVIN L TONG H BOUDAD L MEISTER C PIUZ ATABOUELLE J AARAB M AMIOT R BUFFETAUT EDYKE G HUA S amp LE LOEUFF J 2010 Vertebrateassemblages from the early Late Cretaceous of south-eastern Morocco an overview Journal of African EarthSciences doi101016jjafrearsci200912007 in press

CHARIG A J amp MILNER A C 1997 Baronyx walkeri a fisheating dinosaur from the Wealden of Surrey Bulletin ofthe Natural History Museum Geology Series 53 11ndash70

CHURCHER C S amp DE IULIIS G 2001 A new species ofProtopterus and a revision of Ceratodus humei (DipnoiCeratodontiformes) from the Mut Formation of easternDakhleh Oasis Western Desert of Egypt Journal ofPaleontology 44 305ndash23

CHURCHER C S DE IULIS G amp KLEINDIENST M R2006 A new genus for the dipnoan species Ceratodustuberculatus Tabaste 1963 Geodiversitas 28 635ndash47

COLIN J P EL ALBANI A FURSICH F T MARTIN-CLOSAS C MAZIN J M amp BILLON-BRUYAT J P 2004Le gisement ldquoPurbeckienrdquo de vertebres de Cherves-de-Cognac Charente (SW France) nouvelles donneesbiostratigraphiques Comptes Rendus Palevol 3 9ndash16

CUNY G OUAJA M SRARFI D SCHMITZ L BUFFETAUTE amp BENTON M J 2004 Fossil sharks from the Early



Cretaceous of Tunisia Revue de Paleobiologie specialvolume 9 127ndash42

DALLA VECCHIA F M 1995 Second record of a site withdinosaur skeletal remains in Libya (northern Africa)Natura Nascosta 11 16ndash19

DE CARVALHO M S S amp MAISEY J G 2008 Newoccurrence of Mawsonia (Sarcopterygii Actinistia)from the Early Cretaceous of the Sanfranciscana BasinMinas Gerais southeastern Brazil In Fishes and theBreak-up of Pangaea (eds L Cavin A Longbottom ampM Richter) pp 109ndash44 Geological Society of LondonSpecial Publication no 295

DESIO A 1950 Bibliografia geologica italiana dellrsquoAfricasino al 1948 incluso Collezione scientifica e document-aria dellrsquoAfrica italiana 15 1ndash83

DUFFIN C J 2001 The hybodont shark PriohybodusdrsquoErasmo 1960 (Early Cretaceous northern Africa)Zoological Journal of the Linnean Society 133 303ndash8

EL-ZOUKI A Y 1980 Stratigraphy and lithofacies ofthe continental clastics (Upper Jurassic and LowerCretaceous) of Jabal Nafusah NW Libya In TheGeology of Libya Vol II (eds M J Salem amp M TBusrewil) pp 393ndash418 London Academic Press

FATMI A N ELIAGOUBI B A amp HAMMUDA O S 1980Stratigraphic nomenclature of the Pre Upper CretaceousMesozoic rocks of Jabal Nafusah NW Libya In SecondSymposium on the Geology of Libya Vol 1 (eds M JSalem amp M T Busrewil) pp 57ndash66 London AcademicPress

HAMMUDA O S 1969 Jurassic and Lower Cretaceous rocksof Central Jabal Nefusa Northewestern Libya TripoliLibya The Petroleum Exploration Society of Libya79 pp

KILIAN C 1931 Des principaux complexes continentaux duSahara Comptes-rendus sommaires des seances de laSociete geologique de France 9 109ndash11

KRIWET J amp SCHMITZ L 2005 New insight into thedistribution and palaeobiology of the pycnodont fishGyrodus Acta Palaeontologica Polonica 50 49ndash56

DE LAPPARENT A F 1951 Decouverte de Dinosauriensassocies a une faune de Reptiles et de Poissons dans leCretace inferieur de lrsquoExtreme Sud tunisien Comptes-Rendus de lrsquoAcademie des Sciences de Paris 2321430ndash2

DE LAPPARENT A F 1960 Les Dinosauriens du ldquoContin-ental Intercalairerdquo du Sahara central Memoires de laSociete geologique de France (Nouvelle Serie) 88A1ndash57

LE LOEUFF J BUFFETAUT E amp MERSER C 1996Decouverte drsquoun dinosaure tithonien dans la region deCognac (Charente France) Geologie de la France 279ndash81

MARTIN M 1984 Deux Lepidosirenidae (Dipnoi) cretacesdu Sahara Protopterus humei (Priem) et Protopterusprotopteroides (Tabaste) Palaontologische Zeitschrift58 265ndash77

MILNER A C 2003 Fish-eating theropods a short reviewof the systematics biology and palaeobiogeography ofspinosaurs Actas de las II Jornadas Internacionalessobre Paleontologıa de Dinosaurios y su Entorno(2001) (eds P Huerta Hurtado amp F Torcida Fernandez-Baldor) pp 129ndash38 Salas de Los Infantes ColectivoArqueologicondashPaleontologico de Salas

NOVOVIC T 1977 Sheet Nalut (NI 32ndash4) Geological Mapof Libya scale 1250 000 Explanatory booklet TripoliIndustrial Research Centre 68 pp

OrsquoLEARY M A ROBERTS E M HEAD J J SISSOKOF amp BOUARE M L 2004 Titanosaurian (Dinosauria

Sauropoda) remains from the ldquoContinental intercalairerdquoof Mali Journal of Vertebrate Paleontology 24 923ndash30

OUAJA M FERRY S BARALE G amp SRARFI D 2002Facies de depot du Jurassique et du Cretace du bassinde Tataouine (Sud-Est de la Tunisie) Livret guide delrsquoexcursion organisee par le Service Geologique deTunisie et lrsquoAssociation des Sedimentologistes Francaispublication de lrsquoOffice National de Mines 99 pp

PEREA D UBILLA M ROJAS A amp GOSO C A 2001 TheWest Gondwanan occurrence of the hybodontid sharkPriohybodus and the Late JurassicndashEarly Cretaceous ageof the Tacuarembo Formation Uruguay Palaeontology44 1227ndash35

PERVINQUIERE L 1912 Sur la geologie de lrsquoextreme sud-tunisien et de la Tripolitaine Bulletin de la Societegeologique de France 4 143ndash93

PEYBERNES B VILA J M SOUQUET P CHARRIEREA Ben YOUSSEF M ZARBOUT M amp CALZADA S1996 Trois gisements de brachiopodes dans le Cretaceinferieur tunisien Batalleria 6 45ndash58

RAUHUT O W M amp LOPEZ-ARBARELLO A 2009Considerations on the age of the Tiouaren Formation(Iullemmeden Basin NigerAfrica) Implications forGondwanan Mesozoic terrestrial vertebrate faunas Pa-laeogeography Palaeoclimatology Palaeoecology 271259ndash67

REES J amp UNDERWOOD C J 2008 Hybodont sharks ofthe English Bathonian and Callovian (Middle Jurassic)Palaeontology 51 117ndash47

RICH T H V MOLNAR R E amp RICH P V 1983 Fossilvertebrates from the Late Jurassic of Early CretaceousKirkwood Formation Algoa Basin southern AfricaTransactions of the Geological Society of South Africa86 281ndash91

ROBAUX A CHOUBERT G FLANDRIN J amp DROSDOFF A1942 Notice sur la carte geologique et hydrogeologiqueprovisoire de la Tunisie Feuilles Dehibat et Djenein au1200 000e Direction des Travaux Publics de Tunisie57 pp

ROYO-TORES R COBOS A amp ALCALA L 2006 A GiantEuropean Dinosaur and a New Sauropod Clade Science314 1925ndash7

RUBINO J L LAFONT F METAIS E BLANPIED CFERRY S OUAJA M amp SBETA A 2009 Large scaletidal dunes complex of Aptian ()ndashAlbian age alongNafusah cliff LibyandashTunisia 12eme congres Francaisde Sedimentologie ndash Rennes 2009 Livres des resumesPubl ASF 64 p 289

SANZ J L BUSCALIONI A D CASANOVAS M Lamp SANTAFE J V 1987 Dinosaurios del CretacicoInferior de Galve (Teruel Espana) Estudios GeologicosVolumen Extraordinario Galve-Tremp 45ndash64

SASSI P 1942 Sui Fossili di Due Giascinentie Wealdianidella Tripolitania Annali Museo Libico Storia Natu-rale 3

SERENO P C WILSON J A LARSSON H C E DUTHEILD B amp SUES H-D 1994 Early Cretaceous dinosaursfrom the Sahara Science 266 267ndash71

SERENO P C BECK A L DUTHEIL D B GADO BLARSSON H C E LYON G H MARCOT J DRAUHUT O W M SADLEIR R W SIDOR C AVARRICCHIO D J WILSON G P amp WILSON J A 1998A long-snouted predatory dinosaur from Africa and theevolution of the spinosaurids Science 282 1298ndash1302

SERENO P C BECK A L DUTHEIL D B LARSSON H CE LYON G H MOUSSA B SADLEIR R W SIDORC A VARRICCHIO D J WILSON G P amp WILSONJ A 1999 Cretaceous sauropods from the Sahara and



the uneven rate of skeletal evolution among dinosaursScience 286 1342ndash7

SERENO P C LARSSON H C E SIDOR C A amp GADOB 2001 The giant crocodyliform Sarcosuchus from theCretaceous of Africa Science 294 1516ndash19

SMITH J B amp DALLA VECCHIA F M 2006 An abelisaurid(Dinosauria Theropoda) tooth from the Lower Creta-ceous Chicla formation of Libya Journal of AfricanEarth Sciences 46 240ndash4

SMITH J B amp LAMANNA M C 2009 Abelisauroiddinosaurs from the Early Cretaceous of Libya Geo-logical Society of America Abstracts with Programs41 242

SMITH J B LAMANNA M C LACOVARA K J DODSONP SMITH J R POOLE K J GIEGENGACK R amp ATTIAY 2001 A giant sauropod dinosaur from an UpperCretaceous Mangrove deposit in Egypt Science 2921704ndash6

SMITH J TSHAKREEN S RASMUSSEN S amp LAMANNAM 2006 New dinosaur discoveries from the EarlyCretaceous of Libya Journal of Vertebrate Paleontology26(3) 126

SRARFI D OUAJA M BUFFETAUT E CUNY G BARALEG FERRY S amp FARA E 2004 Position stratigraphiquedes niveaux a vertebres du Mesozoıque du Sud-Est dela Tunisie Notes du Service Geologique de Tunisie 725ndash16

STROMER E 1936 Ergebnisse der Forschungsreisen ProfE Stromers in den Wusten Agyptens VII Baharije-Kessel und -Stufe mit deren Fauna und Flora Eine

erganzende Zusammenfassung Abhandlungen der Bay-erischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung n f 33 1ndash102

TABASTE N 1963 Etude de restes de poissons du Cretacesaharien Memoire IFAN 68 melanges ichthyologiques437ndash85

TAQUET P 1976 Geologie et paleontologie du gise-ment de Gadoufaoua (Aptien du Niger) Cahiers dePaleontologie du C N R S 191 pp

TAQUET P 1984 Une curieuse specialisation du crane decertains dinosaures carnivores du Cretace le museaulong et etroit des Spinosaurides Comptes Rendus delrsquoAcademie des Sciences de Paris II 299 217ndash22

TAQUET P amp RUSSELL D A 1998 New data on spinosauriddinosaur from the Early Cretaceous of the SaharaComptes Rendus de lrsquoAcademie des Sciences de Paris327 347ndash53

TEKBALI A O 2000 Pollen and spores from SubsurfaceAlbian of the Kiklah Formation NW Libya AbstractInternational Workshop on North African Micropale-ontology for petroleum exploration University CollegeLondon 21ndash25 August 2000 p 70

WENZ S 1981 Un coelacanthe geant Mawsonia la-vocati Tabaste de lrsquoAlbienndashbase du Cenomanien duSud marocain Annales de Paleontologie (Vertebres)67 1ndash20

WERNER C 1994 Die kontinentale Wirbeltierfauna aus derunteren Oberkreide des Sudan (Wadi Milk Formation)Berliner geowissenschaftliche Abhandlungen (E) 13221ndash49



lsquo1943 Tunisian eventsrsquo which prevented the authorsfrom correcting the proofs The so-called events arerelated to some episodes of the Second World Warin North Africa when German and Italian troopsinvaded the French protectorate of Tunisia at the endof 1942 before they had to leave the country in 1943following the battle of El Alamein Their defeat led tothe departure of Italians from Libya and the closureof their geological institutions in Africa Howeverwe could not find a single original reference in theAnnali del Museo Libico di Storia Naturale about thesepurported discoveries of fishes and Desio (1950) in hisbibliography of the geology of Libya does not mentionEarly Cretaceous (lsquoWealdenrsquo) fossil fishes found byItalian geologists It thus seems likely that Robaux et alwould have heard about discoveries of fossil fishes inTripolitania by Italian colleagues which were neverpublished and soon forgotten following the collapseof Italian colonies in North Africa An alternativepossibility is that Robaux et al mistakenly mentionedfishes when referring to Sassirsquos recent (1942) reportof Wealden plants in the Early Cretaceous of LibyaAlthough a forgotten publication in some obscurejournal cannot be excluded we suspect that one ofour hypotheses is correct Lapparent was thus the firstauthor to report Early Cretaceous vertebrates fromLibya in 1960 he mentioned several sites discoveredby him along the Dahar escarpment in South Tunisiaand Tripolitania (northwestern Libya) during a fieldtrip in 1951 on the tracks of the French geologistPervinquieres who was the first in 1912 to describefossil vertebrates from the same horizons in southernTunisia In Libya Lapparent mentions several localitiesin Nalut Giado Iefren and Garian The Giado localityyielded a theropod tooth and a bone referred by himto Elaphrosaurus iguidiensis Other sites providedfishes and crocodiles only and were not described indetail by Lapparent Later Burollet amp Manderscheid(1965) mentioned teeth and jaws of crocodiles fromthe Cabao Formation Hammuda mentioned in 1969 acrocodile tooth found at Wadi al-Mezaida near GiadoThe Libyan geologist El-Zouki (1980) later figuredvarious teeth and scales referred to fishes turtles andcrocodiles as well as a dinosaur vertebra from the CabaoFormation in a sand quarry at Jannawan near GiadoAlthough El-Zouki refers this material to Spinosaurus() the published plate (a vertebral centrum in ventralview) does not allow any precise assessment Morerecently Smith amp Dalla Vecchia (2006) describeda possible abelisaurid tooth collected by an Italianamateur palaeontologist near Nalut (see also DallaVecchia 1995 for a preliminary description of thistooth) The same locality has yielded abundant materialof the hybodont shark Priohybodus described by Duffinin 2001 This material has been found according toDuffin (2001) and Smith amp Dalla Vecchia (2006) in theKiklah Formation which overlies the Cabao Formationbut our own investigations in this area suggest thatthis material comes rather from the Cabao FormationSmith et al (2006) and Smith amp Lamanna (2009) in

Figure 1 Location map

two abstracts briefly reported more discoveries in theCabao and Kiklah formations including abelisauroidbones a possible titanosauriform along with sharkturtle lungfish and crocodyliform remains

3 Geology

Several sand quarries located along the northernborder of the Jabal Nafusah have yielded vertebrateremains The material described here was found in threequarries Tendamirah Jawsh and Jannawan (Fig 1)These are all opened in the Cabao Formation acontinental to shallow marine unit (estuarine with tidalinfluences on depositional settings cf Rubino et al2009) first described by Burollet in 1963 (Burollet1963ab) near the village of Giado At this localityit directly overlies the marine Late Jurassic ShakshukFormation and underlies the Kiklah Formation (Fig 2)a possible lateral equivalent of the Oum ed Diab andChenini members of the Aın Guettar Formation ofsouthern Tunisia considered as Late Aptian to Albianin age (see Ouaja et al 2002) Since that time thisstratigraphic succession has been adopted by all authorsworking in Djebal Nefusah (Hammuda 1969 El-Zouki 1980 Fatmi Eliagoubi amp Hammuda 1980)but eight sedimentological sections logged betweenthe Giado locality and the Tunisian border clearlydemonstrate that an intermediate sandy and shalyinterval appears and develops between the Shakshukand Cabao formations in this area (Fig 3) This seriesup to 120 m thick is topped by a distinct memberof lagoonal green shales and dolomites that clearlycorresponds to the Douiret shale first described insouthern Tunisia by Busson in 1967 and ranged morerecently in the upper Douiret Formation by Ouajaet al (2002) This additional interval intercalatedbetween the Shakshuk and the Cabao formations isherein mentioned for the first time in Libya andshould be considered as the lateral equivalent ofthe Tunisian Boulouha and Douiret formations (see























4c Dinosaurs


















6 Conclusions






References





































































































4c Dinosaurs


















6 Conclusions






References






























































































6 Conclusions






References



















































































References














































































