From the Upper Cretaceous of Valencia, Spain

ORYCTOS,VoI.1 : l2I - 126,Octobre 1998

FIRSTDATA ON THE HADROSAURIDDINOSAURS (ORNITHISCHIA, DINOSAURH) FROMTHE UPPERCRETACEOUS OF VALENCIA,SPAIN.

12, Julio COMPANY Angel GALOBART 3 and Rodrigo GAETE 3

' Colegio Universitario C.E.U. San Pablo. Avenida seminario s/n,46113 Moncada, Valencia, Espafla. 'Universidad de Valencia. Departamentode Geologîa,46100 Burjassot, Valencia, Espafla. 3 Institut de Paleontologia "M. Crusafont". Escola Industrial 23,08201, Sabadell,Espafla.

Abstract : Recently,remains of dinosaursassigned to the family Hadrosauridaehave beenrecovered from the Upper Cretaceouslocality of La Solana, Spain. The material presentedhere consists of a lower jaw and severalfragmen- tary isolated teeth. Diagnostic featuresof bonesplace the hadrosaursfrom Spain closer to the hadrosauriddinosaurs from southernFrance than to Tblmatosaurustranssylvanicus from the Maastrichtian of Romania.

Keywords.- Upper Cretaceous,hadrosauridae, dentary, La Solana locality, Spain.

Premièresdonnées sur lesdinosaures hadrosauridés (Dinosauria, Ornithischia) du Crétacésupérieur de Valence,Espagne.

Résumé : Des restesde dinosauresdu Crétacé supérieurde Valence (Espagne)sont rapportésà la famille des Hadrosauridae.Le matériel comprend un fragment de dentaire et plusieurs dents isolées.les caracterfstiquesde ces os les rapprochentde ceux du Maastrichtien du sud-ouestde la Francel ils sont clairement différents de ceux de Telmatosaurustranssylvanicus de Roumanie. (traduit par la rédaction)

Mots clés.- Crétacé supérieu4 dinosaureshadrosauridés, dentaire, La Solana, Espagne.

INTRODUCTION Other specimens are referred to Hadrosauridae indet. (Laurent, 1996; Le Læuff et al. 1993; Different localities in Europe have provided Wellnhofer, 1994; Mulder et al., 1997) remains of hadrosaurid dinosaurs (fig. 1) (Le Læuff et al., 1993). At present, some of these specimens have been assigned to three different genera: GEOGRAPHIC AND GEOLOGIC SETTING Telmatosattrus, Orthomerus and Pararhabdodon, although several authors consider a possible synony- The vertebrate locality of La Solana is located my between Telmatosaurus and Orthomerus (Paris near the village of Carlet, in Tous County, about 40 & Taquet, 1973; Brinkmann, 1984; Mulder 1984). km. to the SW of Valencia, Spain (fig. 1). The most complete specimens belong to The outcrop is placed in the southeasternmargin Telmatosaurus transsylvanicus from the Hateg of the Iberian Chain and comprises the uppermost Region in Romania (Nopcsa, 1900, 1903; member of the continental sediments of the Weishampel et al., 199I, 1993). Orthomerus dolloi, Limestones and Marls of the Sierra Perenchiza from the Maastrichtian of Belgium (Buffetaut et al., Formation (Campanian-Maastrichtian : Gutiérrez et 1985; Brinkmann, 1988), and Pararhabdodon iso- al., 1984). In the area studied, these beds are folded nensis (Casanovas et al., 1993, 1997; Laurent, in a syncline with nearly continuous exposuresalong 1996), from the Tremp Basin (Spain) and southern a E-W trend. Bones were collected from continental France, ate two other hadrosaurid dinosaurs,but they beds arranged in a sequence of fine-grained sand- are actually known only by scarce cranial and post- stones, silty marls and dark clays. This unit is over- cranial elements. lain by fluvial Paleogenedeposits.

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Fig.l - Geographic distribution of hadrosaur material. A: location map of Europe showing the main Late Cretaceousdinosaur localities where remains of hadrosaurs have been collected. 1: Hateg Basin (Transylvania, Romania );2: Bakhtchisarar (Crimea, Ucrania); 3: Localities of Le Jadet,Auzas, Tricouté and Lestaillats (Haute Garonne, France); 4: Mérigon (Ariège, France); 5: Le Bexen (Aude, France); 6: Maastricht area (Limburg, Belgium and Netherlands); 7: Els Nerets and Sant Romà d'Abella (Tremp Basin, Spain); 8: La Solana (Valencia, Spain); 9: BadAdelholzen (Bavaria, Germany). B: location of La Solana locality, near the village of Carlet, in the province ofValencia (Spain).

Fig.1 - Distribution géographiquedes hadrosauridés.A: carte d'Europe montrant les principaux gisementsà hadrosauridésdu CréIacésupérieur. B: localisation du gisementde La Solana,près du village de Carlet, province deValencia(Espagne)

A preliminary list of the findings includes plant 220I) and two fragmented maxillary teeth (MGUV microfossils and carbonaceousplant remains, fresh- 2232, MGUV 2233). These teeth vary considerably water gastropods, ostracods and bivalves, abundant in dimensions, so we believe that they belong to indi- fish scales, fragments of bones of chelonians and viduals of different size. small amphibians, crocodilian teeth, and fragmentary remains of hadrosaurid dinosaurs. The fossil assemblage and the sedimentological features sug- DESCRIPTION gest a lacustrine or a swamp environment. Dentary. MGUV 2200 is a I82 mm long, left dentary which preserves several tooth families and MATERIAL part of the bony plate that originally covered them (fig. 2). This specimen is not complete, as the rostral The material collected consists of fragmentary end is not preserved. cranial and postcranial elements (an incomplete left The body of the dentary is rectangular, with the dentary, severalisolated teeth, fragmentary ribs, seve- dorsal margin almost straight and the ventral side ral fragmented vertebral centra, the proximal end of slightly convex. The coronoid process,highly develo- the left humerus and a fuagment of the right femoral ped, arises from the caudal end of the laterul surface shaft). The specimensdescribed here are a left denta- of the dentary. It extends outward and upward, per- ry (MGUV 2200), a single dentary tooth (MGUV pendicularly to the main body of the dentary. r22 COMPANY, GALOBART & GAETE - SPANISHHADROSAURIDS

The lateral surface of the dentary is strongly The crown is lanceolate in lingual view and is convex in its posterior portion, where the base of the almost straight in mesiodistal view. The crown flanks coronoid process is formed, and concave in its ante- are flat, and the edges bear small marginal denticles rior part. It presents five small foramina that are not supported by ridges. This feature differs clearly directed backwards. from that of T. transsylvanicus (Weishampel et al., Over the medial dentary face, 18 complete or r99r, 1993). fragmentary teeth are preserved, affanged in 12 verti- MGUV 2232 and MGUV 2233 aretwo large, cal rows and partially covered by a thin bony plate hadrosaurmaxillary teeth.MGUV 2232 includesthe (tooth description below). About 17 dental alveoli are apexend of a crown and MGUY 2233 (fig. 4) is an present too, although the rostral ones are incomplete- almost completetooth with a well-preservedcrown ly preserved. Consequently, there are approximately (maximutrnwidth , II mm; length, 32 mm) and a bro- 29 tooth positions. The alveoli, about 3-5 mm wide, ken root. Teeth show no sign of wear. Both teeth are bend forwards and decrease in size rostrally. Along squareto subrhomboidalin cross section.In mesio- the medial side of the dentary, below the alveoli, a distal view, the crown of each individual tooth is row of foramina, partially preserved, opens through strongly curvedlingually. The crown is elongate and the bony plate. A large cavity, the mandibular fossa, only its buccal face is enameled.It presentsa high opens posteriorly in the caudal side of the dentary. longitudinal carina that is offset distally, and two This cavity extends forwards as an elliptical foramen shortsecondary ridges at the baseof the mesialregion within the dentary: the supra-Meckelian foramen. The of the crown. Between the median carina and the greater axis of this opening is oblique, while it is ver- marginalrims, the crown flanks arecurved. The mar- tical in klmatosaurus transsylvanicus (Le Læuff et gins of the crown are elevatedand display smooth al., 1993). On the other hand, the mandibular fossa papillations. continues downwards as a naffow groove, the Mecke- lian canal, that runs forwards through the ventral margin of the dentary. The angular and splenial, which form the floor of the Meckelian canal, are missing.

Dentition MGUV 220I is an unworn small-sized hadrosaur dentary tooth (8 mm wide) that includes one-third of the apex end of a crown. It presentsthe samefeatures than those of the teeth preserved in MGUV 2200 (fig. 3). The crown is enameled only on the lingual face and is ornamented by a sharp median carina and by a faint secondary ridge, mesially placed.

Fig.2 - Left dentary of Hadrosauridae indet. (MGUY 2200) from La Solana locality. A: medial view. B: lateral view. Abbreviations: Cp, coronoid process;Db, dental battery; A1, alveoli; Mc, Mec- kelian canal; Mt mental foramina. Scalebar = 50 mm.

Fig.2 - Dentaire gauche MGUV 2200 du gisement de La Solana. A: vue médiale. B: vue lat&ale. Abréviations: Cp, processuscoronoïde; Db, batte- rie dental; Al, alvéoles;Mc: canal de Meckel; Mf, foramensmentonniers. Barre d'échelle = 50 film. 123 ORYCTOS,Vol. 1, 1998

Fig.3 - Comparison of dentary teeth of Late Cretaceous :::::i1r-* hadrosaursfrom Europe.A: lingual view of dentary tooth of Hadrosauridae indet. (MDE-Ma1-01) from Tricouté (Fr), (Redrawn from Laurent, 1996). B: lingual view of left dentary tooth of Hadrosauridaeindet. (MGUY 2200; in situ rooth) from La Solana(Sp) with an enlargementof mesial denticles. C: lingual view of dentary tooth of T. transsylvanicus from HategBasin (Ro), (AfterV/eishampelet a\.,1993). Scalebar for A-D = 10 mm.

Fig.3 - Dents dentaires d'hadrosauridésdes gisements d'Europe. A: dent dentaire d'Hadrosauridae indet. (MDE- Ma1-01) de Tricouté(France). Vue linguale (d'aprèsLaurent, 1996). B: dent dentaire,Hadrosauridae indet (MGUY 2200) du gisementde La Solana(Espagne) avec un agrandissement des denticulesdistaux (vue linguale). C: dent dentaire de T. B A ç transsylvanicusdu Bassin du Hateg Basin, Romania (d'après î-ælq;l reffi ffi Weishampelet aL.,1993).Barres d'échelle = 10 mm.

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Fig.4 - Right maxillary tooth of Hadrosauridaeindet. (MGUV Fig. 4 - Dent maxillaire d'Hadrosauridaeindet (MGUY 2233) 2233) from La Sol anaLocality. A: distal view; B: buccal view du gisementde La Solanaavec un agrandissementdes denti- with an enlargementof marginal papillations;C: mesial view; cules marginaux.Barre d'échelle = 10 mm. D: occlusalview. Scalebar = 10 mm.

These specimens differ from the European DISCUSSION hadrosaurid dinosaur T. transsylvanicus in dental morphology: dentary teeth of Telmatosaurus are MGUV 2200 and MGUV 220I. The general recurved distally, and mesial denticles aîe supported morphology of the lower jaw, the large number of by secondary ridges (Weishampel et al., 1993: text- alveoli, the presence of dental batteries of packed fig. 3G). These features afe not present neither in tooth families, the lanceolate crown of teeth with a MGUV 220I nor in situ dentary teeth. large median carina and the presence of small margi- On the other hand, dentition is very similar to nal denticles are typical characters of the family the specimenMDE-Ma1-01 describedand figured by Hadrosauridae (Lull and Wright, 1942; Weishampel, Laurent (1996) -collected from Tricouté,France- and 1984; Weishampel and Horner, 1990). classified as Hadrosauridaeindet. (Fig. 3). r24 COMPANY,GALOBART & GAETE- SPANISHHADROSAURIDS

Likewise the positioning and orientation of the REFBRENCES supra-Meckelian forameû, the disposition of the den- BRINKMANN, W. 1984. Erster nachweis eines Hadrosauriers (Ornithischia) (Maastrichtium) tal alveoli and the general morphology of the dentary aus dem unteren Garumnium des Beckens von Tremp (Provinz Lérida, Spanien). are characters that place the hadrosaurids from La PalciontologischesZeitschrift, 58 : 295-305. Solana locality closer to those from southern France 1988. Zur fundgeschichteund Systematikder (Laurent, 1996; Le Læuff et al. 1993) than to T. ornithopoden (Ornithischia, Reptilia) aus der Ober-Kreide von transsylvanicus from the Maastrichtian of Romania. Europa.Documenta Naturqe, 45 :I-I5l . The type species of Pararhabdodon and Ortho- BUFFETAUI E.; MEIJER A.W.F.; TAQUET P. & WOUTERS, c. 1985. New merus are based on fragmentary specimens that do remains of hadrosaurian dinosaurs (Reptilia, Ornithischia) from the Maastrichtian of Dutch and Belgiam not include teeth, so it is not possible to refer these Limburg. Revuede Paléobiologie,4 : 65-70. specimensto those genera. CASANOVAS-CLADELLAS, M.L.; SANTAFÉ-LLOPIS, J.V. &. MGUV 2232 and MGUV 2233. Both isolated ISIDRO-LLORENS, A. 1993.Pararhabdodon isonense n. gen. and fragmentary maxillary teeth present the morpho- n. sp. (Dinosauria). Estudio morfol6gico, radio-tomogrâfico y logical characteristics of hadrosaurid dentition. The consideracionesbiomecânicas . Paleontologîay evoluciô, 26-27 : I2t-13 1. shape of the enameled surface and the absence of numerous ridges on the crown distinguish clearly PEREDA-SUBERBIOLA. X. 1997. Nouveaux restes de these specimens from other Late Cretaceousornitho- Pararhabdodon isonensis(Dinosauria, Hadrosauridae)et synthè- pods like Rhabdodon (see Brinkmann, 1988; Wei- se de I'ensembledu matériel découvertdans le Crétacésupérieur shampel et al., I99I). de Catalogne. Second European Workshop of Vertebrate The maxillary teeth from La Solana can be dis- Palaeontology,Espéraza - Quillan, Abstracts. GUTIÉRREZ, tinguished from those of T. transsylvanicus by the G.; F,LIZAGA, E.; GOy, J.L.; NIETO, M. & ROBLES, F. 1984.Mapa Geoldgico de la Provincia de Valencia. absenceof denticles on the margins of the crown and Escala 1:200.000.Diputacidn Provincial de Valencia, Universi- by the presence of upturned and papillated (not den- dad de Valencia,IGME. Valencia.61 pp. ticulated) marginal rims. LAURENT, Y. 1996. Les hadrosaures du Maastrichtien du Sud- Therefore, these remains belong to an hadrosau- Ouestde la France.Mémoire de D.E.A., Université de Montpel- rid dinosaur different from klmatosaurus transsylva- lier II : 1-35 (unpublished). LE LGUFF, nicus and similar to the specimenscollected in sou- J.; BUFFETAUT, E.; MARTIN, M.; MARTIN, V. &. TONG, H. 1993; Découverte d'Hadrosauridae (Dinosauria, thern France. Until a more complete and careful ana- Ornithischia) dans le Maastrichtien des Corbières (Aude, lysis of postcranial elements, they have been provi- France). ComptesRendus de l'Académie des Sciencesde Paris, sionally attributed to Hadrosauridae indet. 316 : 1023-1029. LULL, R.S. 8. WRIGHT, N.V. 1942. Hadrosauriandinosaurs of North America. Geological SocieQ of America Special Papers, ACKNOWLEDGMENTS 40 : I-242. MULDER, E.V/.A. 1984. Resten van Telmatosaurus(Ornithischia, Hadrosauridae) uit het. Boven-Krijt van Zuid-Limburg-Grond- The authors would like to thank Dr. Miquel De boor en Hamer 314: 108-115. Renzi (Universidad de Valencia) for his comments on KUYPERS,M.M.M.; JAGT J.V/.M. & PEETERS an earlier version of this paper. We specially appre- H. H.G. 1991. A new late Maastrichtian hadrosaurid dinosaur ciate the valuable suggestionsmade by reviewers Dr. record from northeast Belgium. Neues Jahrbuch fiir Geologie David V/eishampel (John Hopkins University, Balti- und Palciontologie,Monatshefte : 339-341. NOPCSA, F. 1900. Dinosaurierresteaus Siebenbûrgen.I:(Schâdel more), Dr. Jean Le Loeuff and Yves Laurent (Musée 'We von Limnosaurus transsylvanicus nov. gen.et spec.). des Dinosaures d'Espéraza). also acknowledge Denkschriften der kôniglichen Akademie der Wissenschaften, the technical assistanceof Francisco Periago (Uni- Wien,68: I-37 . versidad de Valencia). -1903. Telmatosaurus,a new name for the dinosaur Financial support for the excavation tasks was LimnosaurLts.Geolo gical Magazine, 2 : 24I-295. PARIS, provided by the Direccid General de Patrimonio J.P. &. TAQUEI P. 1973. Découverte d'un fragment de dentaire d'Hadrosaurien (Reptile Dinosaurien) dans le Crétacé Artfstico (Generalitat Valenciana). Additional finan- supérieur des Petites Pyrénées (Haute-Garonne).Bulletin du cial assistancewas provided by Institucid Valenciana Muséum National d'Histoire Naturelle, 130 : 17-27 . d'Estudis i investigacid( I.V.E.I., grantCPE/098). WEISHAMPEL. D.B. 1984. Evolution of Jaw Mechanics in r25 ORYCTOS, Vol. 1, 1998

ornithopod dinosaurs. Advances in Anatomy, Embriology and Cell Biology,ST : 1-110. GRIGORESCU, D. &. NORMAN, D.B. 199T; The dinosaurs of Transylvania: island biogeography in the Late Cretaceous.National Research& exploration ,7 : 68-87. - & HORNER, J. R. 1990. Hadrosauridae,pp. 534- 561. In WEISHAMPEL, D. B.; DODSON, P. and OSMOLS- KA, H. (eds.) The Dinosauria. University of California Press, Berkeley. - NORMAN, D. B. &. GRIGORESCU, D. 1993. Telmatosaurus transsylvanicus of Romania: The most basal hadrosauriddinosaur. Paleontology, 36 : 361-385. WELLNHOFER, P. 1994. Ein dinosaurier (Hadrosauridae) aus der Oberkreide (Maastricht, Helvetikum-zone) des bayerischen Alpenvorlandes.Mitt. Bayer. Staatsslg.Palaônt, Hist. Geol., 34 : 221-238.

Note reçue le 3l-08-1997 acceptéeaprès révision le l5-03-1998

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