The Site of La Cantalera (Teruel): an Exceptional Window on the Vertebrate Biodiversity of the Hauterivian-Barremian Boundary in the Iberian Peninsula

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THE SITE OF LA CANTALERA (TERUEL): AN EXCEPTIONAL WINDOW ON THE VERTEBRATE BIODIVERSITY OF THE HAUTERIVIAN-BARREMIAN BOUNDARY IN THE IBERIAN PENINSULA

Canudo, J. I. 1, Aurell, M. 2, Badiola, A. 1, Blain, H.-A. 3, Gasca, J. M.1, Gómez-Fernández, D. 1, Moreno-Azanza, M. 1, & Ruiz-Omeñaca, J. I. 4 1. Grupo Aragosaurus-IUCA (http://www.aragosaurus.com). Universidad de Zaragoza. 50009 Zaragoza, Spain 2. Estratigrafía. Universidad de Zaragoza. 50009 Zaragoza. Spain 3. Institut Català de Paleoecologia Humana y Evolució Social (Unitat asociada al CSIC). Universitat Rovira i Virgili. 43005 Tarragona. Spain 4. Museo del Jurásico de Asturias (Muja). 33320 Colunga. Asturias. Spain

YACIMIENTO DE TER LA CANTALERA TER Conglomerados y lutitas 1 JM 2 * C3 C2 C3 Fm. Alacón C1 Calizas bioclásticas (c. 50 m)

JI5 3 C2 Fm. Blesa (superior)

JI4 Calizas y margas (c.100 m)

C1 Fm. Blesa (inferior) JI3 4 Río Cantalera CRETÁCICO INFERIOR MIOCENO Lutitas rojas (50-140 m) Geological map of La Cantalera JI2 JUR. JM Fms. Chelva MED- y Pozuel which is located in the northwes- SUP. Calizas oolíticas (c.50 m)

JI5 Fm. Turmiel tern margin of the Oliete sub- JI1 Margas con fósiles (c.50 m) basin, within the large cretacic TR2 JI4 Fm. Barahona 2 TR1 Maestrazgo Basin (Iberian Basin). Calizas bioclásticas (25 m) Fms. Almonacid JI3 y Cerro Pez Geographic setting of La Cantale- 41º00' Margas y calizas (30 m) 00º50' ra: Josa, Teruel province, Spain. Fm. Cuevas JI2 * Labradas JURÁSICO INFERIOR 1. Ornithopod fibula in situ Cortes de Aragón Calizas tableadas (c. 100 m) Fm. Cortes JI1 de Tajuña 2. Fieldwork during a excavation in La Cantalera site YACIMIENTO DE Carbonatos masivos (150-250 m) LA CANTALER*A Josa

3. General view of the La Cantalera outcrops. * TR2 Imón Fm

Dolomías tableadas (30 m) 4. Partial caudal serie of an undetermined ornithopod dinosaur alera

Río Cant TR1 Keuper facies N 400 m 1 2 km Lutitas y yesos (c.100 m)

La Hoz de la Vieja TRIÁSICO SUPERIOR

The site of La Cantalera is located near the village of Josa 3 (Iberian Range, Teruel, Spain). Geologically, the La Cantalera site is an outcrop of Early Cretaceous clays (“Wealden facies”), specifica- lly clays from the Blesa Formation. The fossiliferous beds compri- se grey clays with abundant vegetal remains, charophytes, ostra- cods and gastropods. Atopochara trivolvis triquetra dates the locality as late Hauterivian-early Barremian in age (subzone triquetra). The site was formed in a palustrine environment with pe- riodic droughts. It is likely to have been a rich and diverse te- rrestrial ecosystem. The highly diverse vertebrate fossils are Blesa Formation: Stratighaphic section and out- croups in La Cantalera area. La Cantalera composed mainly of isolated teeth, disarticulated, complete or fossil site is located in the basal part of this lithostratigraphic fragmented bones, eggshell fragments, and coprolites. To date, 4 unit. 29 different taxa have been identified.

MPZ 97/464. Left upper tooth of the La Cantalera sauropod. Mesial (A), The dinosaurs display exceptional biodiversity. The theropods are lingual (B), distal (C), labial (D), apical (E) and adapical represented by at least 10 tetanurans. There are medium-sized (F) views. The arrows show the incision (i) and the attritional surfaces basal forms included in Spinosauridae and Allosauroidea and a (sa). The labial side is at the bottom in E and F. Scale bar = 10 mm. Canudo et al (2002) great abundance of small or medium-sized maniraptorans attributed to aff. Paronychodon sp., Maniraptora indet. 1, 2 and 3, Dromaeosaurinae indet. 1, 2 and 3, and Velociraptorinae MPZ 2002/926. Polacanthidae indet. Left dentary tooth in lingual (A), labial indet. The sauropods are represented by isolated teeth attri- (B), occlusal (C), mesial (D) and distal (E) buted to Euhelopodidae indet. Shed teeth from ornitho- views. Scale bar 1 mm. Canudo et al. (2004) pods of various sizes are the most abundant dinosaur fossil remains, attributed in large measure to “Hypsilo- phodontidae” indet. and to Iguanodontoidea indet. Less frequent are the dental crowns similar to Ha- drosauridae and small-sized Rhabdodontidae. The thyreophorans are represented by dermal plates and Shed teeth of “Hypsilophodontidae” indet isolated teeth attributed to Polacanthidae indet. THEROPOD TEETH from La Cantalera. a, CAN1-957 Bar- yonychinae indet. b, CAN1- Iguanodontoidea indet. The most abundant crocodylomorph teeth are those 963 Carcharodontosauridae? Right maxillary tooth in of Atoposauridae indet. and Bernissartiidae indet., indet. c, CAN1-979 Dro- labial view, with the root preserved and a developed aff. Chiquitosuchus sp. and Goniopholididae indet. Chelo- maeosaurinae indet. 1. d, wear facet. nians are represented by small-sized peripheral plates included CAN1-1035 aff. Paronycho- (MPZ 97/470) in ?Pleurosternidae There are isolated teeth of at least two diffe- don sp. e, CAN1-1043 Mani- raptora indet. 2. f, rent pterosaurs. Mammals are represented by isolated multitu- MPZ2003/232 Maniraptora ORNITHOPODS are the dinosaur remains most indet. 3. g, CAN1-1031 Mani- abundant within La Cantalera fossil assemblage. berculate teeth. The fossils have been assigned to at least three raptora indet. 1. h, CAN1- Extremely abundant isolated teeth and little taxa: a pinheirodontid taxon, Cantalera abadi; a representative 1001 Dromaeosaurinae indet. postcranial remains represent this fossil assem- of the eobaatarids Eobaatar; a taxon described as Plagiaulacidae 2. i, CAN1-1058 Dromaeo- blage. The postcranial remains are usually frag- saurinae indet. 3. j, CAN1- mentary and disarticulated elements, such as or Eobaataridae gen. et sp. indet.; and another described as Pla- isolated vertebral centra, except a partially arti- giaulacida indet. Lissamphibians are represented by humeri and 1029 Velociraptorinae indet. culated caudal serie, which belongs to an unde- Scale bar: a, b, c 5mm; d termined ornithopod dinosaur. Several verte- dentaries of Albanerpetonidae, as well as a humerus and frag- 0,5mm; e 2mm; f 5mm; g, brae with amphicelic centra and hexagonal ments of tibiofibula of a discoglossid frog (Alytidae). Lacertids h, i, j 1mm. cross-section, typical of Ornithopoda, form this are represented by dentaries, indeterminate tooth-bearing frag- Gasca et al. (2009) semi-articulated caudal serie. However the dental record is more diagnostic and allows dis- ments, and one fragment of vertebra of Paramacellodidae. tinguish several morphotypes that can be assigned to different taxa: Iguanodontoidea indet., “Hypsilophodontidae” indet., ?Hadrosauroidea 1 indet., ?Rhabdodontidae indet. Ornithopod teeth. 1-3, 5: Iguanodontoidea The ornithopod teeth are mostly terminally indet. Dentary tooth in mesial (1) and lingual (5) freshwater gastropod resorbed teeth. It has been collected around views. Maxillary tooth in lingual (2) and labial 1 2 3 400 specimens. The majority are shed teeth (3). 4: “Hypsilophodontidae” indet. Dentary and only a few are functional crowns or replace- tooth. Scale bar 1-5 = 10 mm Other fossil vertebrates.Crocodile (1, 2) and pterosaur teeth (3). 1: Bernissartiidae indet. 2: Atoposauridae indet. ment teeth with the enamel preserved. The exceptional accumulation of shed teeth support 2 3 the hypothesis that La Cantalera area was a place where the phytophagous dinosaurs lost Acknowledgements. Financial support was provided by the Mi-

resorbed teeth from their dental batteries Cantalera abadi Badiola et al. (2008). A: left nisterio de Educación y Ciencia (research project CGL2007 ⁄ during feeding. In the terminally replacement Mammals M1, holotype FCPT (CAN 1/1609) B: left P4/5, 62469 ⁄ BTE “Controles paleogeográficos y paleoclimáticos en stages more teeth became undefined losing paratype FCPT (CAN 1/934). Scale bar 1 mm. el relevo de las faunas de vertebrados continentales durante el completely the original morphology. As a result intervalo Jurásico Superior – Cretácico Inferior en Iberia y Pa- a third of the ornithopod teeth collection cannot The fossil mammal from La Cantalera have increased the resolution of tagonia”) and the Gobierno de Aragón: Departamento de Cien- be assigned to any dental morphotype. Between European Early Cretaceous multituberculate mammalian biodiversity, cia, Tecnología y Universidad (Financiación de Grupos Consoli- identifiable teeth the 80% have been assigned biostratigraphy and palaeobiogeography: (1) The discovery of a new late dados, 2008) and the Dirección General de Patrimonio Cultural to Iguanodontoidea (216 specimens). Less fre- Hauterivian pinheirodontid taxon demonstrates greater biodiversity and which have supported the prospection fieldwork. quent are the “Hypsilophodontid” morphotypes a wider biostratigraphic distribution for these multituberculates than was (32 specimens). Only a few teeth can be inclu- previously known. (2) The oldest representative of Eobaatar is described ded within other ornithopod morphotypes that in Western Europe, in the Iberian locality of La Cantalera. (3) The taxon resembles hadrosauroid crowns (16 specimens) described as Plagiaulacidae or Eobaataridae in La Cantalera has been and one fragment of a crown shows features also identified in another Early Cretaceous (early Barremian) bed of the similar to small-sized Rhabdodontidae. 4 5 Aragonese branch of the Iberian Ranges, in Galve (Teruel, Spain).