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Home , Ceresiosaurus, Ichthyosaurus, Lariosaurus, Nothosaurus, Paraplacodus, Pistosauroidea

marine from the Iberian Peninsula

Los reptiles marinos del Mesozoico de la Península Ibérica

N. Bardet 1, X. Pereda Suberbiola 2 y J.I. Ruiz Omeñaca 3

1 UMR 5143 du CNRS, Muséum national d’Histoire naturelle, 57 rue Cuvier, C.P. 38, 75005 Paris, . [email protected] 2 Departamento de Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Universidad del País Vasco/EHU, Apdo. 644, 48080 Bilbao, . [email protected] 3 Museo del Jurásico de Asturias, 33328 Colunga, Spain. [email protected]

Abstract: A synthesis of the record of Mesozoic marine reptiles from the Iberian Peninsula (Spain and ) is provided for the first time. Though this fossil record appears poor compared to other European regions, it astonishes by its expanded stratigraphical range, from the Middle to the latest , as well as by its high-level taxonomical diversity, most being represented, i.e. thalattosaurs, sauropterygians, ichthyosaurians, mosasaurids, , crocodyliforms and .

Key words: , , Ichthyosauria, , Crocodyliformes, Testudines, Spain, Portugal.

Resumen : Se presenta por primera vez una síntesis del registro fósil de reptiles marinos mesozoicos en la Península Ibérica (España y Portugal). Aunque dicho registro pueda parecer pobre comparado con otras regiones europeas, es sorprendente por su amplio rango estratigráfico, desde el Triásico medio al Cretácico Superior, así como por su gran diversidad taxonómica a nivel de taxones de alto rango, estando representados la mayoría de los clados, a saber: talatosaurios, sauropterigios, ictiosaurios, mosasáuridos, serpientes, cocodrilos y tortugas.

Palabras clave: Thalattosauria, Sauropterygia, Ichthyosauria, Squamata, Crocodyliformes, Testudines, España, Portugal.

INTRODUCTION GEOGRAPHY AND GEOLOGY (Fig. 1)

The first mentions of the discovery of Mesozoic Triassic – Marine remains are abundant in in the Iberian Peninsula date from the ( facies) and, to a lesser the second half of the XIX th century (Schulz, 1858 for extent, in the Upper Triassic ( facies) of Spain. Spain; Sauvage, 1897-1898 for Portugal), roughly Up to now, they have not been unearthed in the Triassic coinciding with those made in the rest of and of Portugal nor in the Lower Triassic of Spain. All linked to the Industrial Revolution engineering works. outcrops are located in the eastern part of the Iberian Since that time and though good exposures of marine Peninsula, having been found in the Autonomous Mesozoic rocks, the fossil record of Iberian marine Communities of Aragón (Huesca and Teruel provinces), reptiles remains quite poor – both quantitatively and Castilla-La Mancha (Cuenca, Albacete, Guadalajara and qualitatively – as compared to other European countries. Ciudad Real), Cataluña (Barcelona and Tarragona) and Indeed, Lagerstätten outcrops are unknown for Iberian Islas Baleares (Mallorca). The most fossiliferous marine reptiles, which are mainly represented by outcrops are those of Mont-ral-Alcover (Tarragona, isolated and fragmentary specimens (vertebrae and , Fig. 1, n°4), which has yielded the greatest teeth), producing very few determinable infra-familial number of specimens and taxa; Estada (Huesca, Mid- taxa. However, the Iberian record of Mesozoic marine Triassic, n°1) which is the oldest known one; and reptiles astonishes by its expanded stratigraphical range, Manzarena (Teruel, , n°8), which is the only from Middle Triassic to uppermost Cretaceous, and by outcrop of sure age from the Iberian its high-level taxonomical diversity, with most of the Peninsula. major clades being represented. The aim of this paper is to present an overview of the Mesozoic marine reptiles – Jurassic outcrops are more numerous of the Iberian Peninsula, both on a spatiotemporal and a than Triassic ones, being known both in Portugal and systematical perspective. The bibliographic database Spain. They range from the Lower Liassic () used is Quesada et al . (1998) for Spain and Crespo to the Upper Malm (). Lower Jurassic outcrops (2002) for Portugal, complemented by more recent are abundant, especially those of age, a references when necessary. stage worldwidly considered as very poor in marine reptile remains. Upper Jurassic outcrops are also well represented, especially those of age.

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Middle Jurassic marine reptiles are poorly known and represented by remains only.

Contrary to Triassic fossiliferous sites, the Jurassic ones are concentrated on the western part of the Iberian Peninsula, and more especially on the Atlantic Coast of Portugal (Central region) and Spain (Asturian coast). Specimens from Spain come from Asturias, Castilla y León (Soria province), La Rioja, Aragón (Zaragoza and Teruel) and Valencia. Those from Portugal have been unearthed from the Central (Aveiro, Coimbra and Leiria districts), Lisboa (Lisboa district) and Algarve (Faro district) regions. The most productive levels are those of: 1) the Lias of the Asturias coast (Fig. 1, n°16) and of the Central region of Portugal (n°35-38, 40, 41, 43, 44); and 2) the Kimmeridgian of the same areas of Spain FIGURE 1. Map of the main localities with Mesozoic marine reptiles (n°17, 18) and Portugal (n°38, 41, 42, 45). The first in Spain (provinces in brackets) and Portugal (districts in brackets). SPAIN - Triassic : 1: Estada (Huesca); 2: Aiguafreda (Barcelona); 3: marine reptile remains of Iberia were found in these Olesa (Barcelona); 4: Mont-ral-Alcover (Tarragona); 5: Riba de areas (Schulz, 1858; Sauvage, 1897-1898). Santiuste-Tordelrábano (Guadalajara); 6: Rillo de Gallo (Guadalajara); 7: Royuela (Teruel); 8: Manzanera, Torrijas (Teruel); Cretaceous – Marine reptile fossils are rather scarce 9: Valdemeca (Cuenca); 10 : Boniches, Cañete (Cuenca); 11 : Henarejos (Cuenca); 12 : Sóller (Mallorca); 13 : Alcázar de San Juan in the Cretaceous of the Iberian Peninsula and come (Cuidad Real); 14 : Alpera (Albacete); 15 : Bienservida-Villarrodrigo from the Lower Cretaceous ( to , (Albacete). Jurassic : 16 : Villaviciosa (Asturias); 17 : Colunga including Wealden facies) and Upper Cretaceous rocks (Asturias); 18 : Ribadesella (Asturias); 19 : Valgañón (La Rioja); 20 : (, to ). Ólvega (Soria); 21 : Ricla (Zaragoza); 22 : Obón (Teruel); 23 : Buñol (Valencia). Cretaceous : 24 : San Pantaleón de Losa (Burgos), Bóveda (Álava); 25 : Castillo-Lasarte (Álava); 26 : Jáuregui (Álava); 27 : Laño The outcrops are located in the País Vasco (Álava), (Condado de Treviño); 28 : Ortigosa de Cameros (La Rioja); 29 : Galve Castilla y León (Burgos and Condado de Treviño), La (Teruel); 30 : Castellote, Las Parras de Castellote (Teruel); 31 : Alcalá Rioja, Castilla-La Mancha (Cuenca), Aragón (Teruel) de la Selva (Teruel); 32 : Morella, Todolella (Castellón); 33 : Las Hoyas (Cuenca); 34 : Benagéber (Valencia). and Valencia (Castellón and Valencia provinces) in PORTUGAL - Jurassic : 35 : Casal Comba (Aveiro); 36 : Cadima, Spain, and in the Central (Aveiro and Coimbra Murtede (Coimbra); 37 : Alhadas (Coimbra); 38 : Condeixa-a-Nova/ districts), Lisboa (Lisboa district) and Algarve (Faro Miranda do Corvo, Cezimbra (Coimbra); 39 : Pombal (Leiria); 40 : Foz district) regions of Portugal. The most fossiliferous sites da Ribeira, Agua de Madeiros (Leiria); 41 , Praia N.S. da Victoria, Pintanheira, Fervença, Romão (Leiria); 42 : Guimarota (Leiria); 43 : are those of: 1) the latest Cretaceous of the Álava and Alvaiázere (Leiria); 44 : Tomar (Santarém); 45 : Cesareda (Leiria); 46 : Burgos provinces (Fig.1, n°24-26), mainly the Laño Lourinhã, Praia de Santa Cruz (Lisboa); 47 : Vila Franca do Rosário quarry (n°27), and of Aveiro (n°49) and Coimbra (n°51, (Lisboa); 48 : Malhão (Faro). Cretaceous : 49 : Aveiro (Aveiro); 50 : 52) districts; 2) the Cenomanian of Lisboa (n°53); and Chousa do Fidalgo (Aveiro); 51 : Viso (Coimbra); 52 : Taveiro (Coimbra); 53 : Pendao/Figueira, Cacém, São Pedro do Estoril 3) the Wealden of Valencia (n°34). (Lisboa); 54 : Alcantara (Faro).

SYSTEMATICS Placodonts are mainly known by characteristic rounded, low, crushing teeth (Pinna, 1990 in Quesada et Most Mesozoic marine reptile groups are al., 1998; Rieppel, 2000). Two genera are known: the represented in the fossil record of the Iberian Peninsula: placodontoid in the Muschelkalk/ thalattosaurs, sauropterygians, ichthyopterygians, Keuper? of Henajeros (n°11) and the cyamodontoid squamates, crocodyliforms and chelonians. ? in the Ladinian of Mont-ral-Alcover (n°4). Cyamodontoids are also represented by Thalattosauria – The only mention of this Triassic placochelyids in the Muschelkalk of Bienservida- group has been made in the Ladinian of Mont-ral- Villarrodrigo (n°15) and in the Muschelkalk/Keuper? of Alcover (Fig. 1, n°4) (Rieppel y Hagdorn, 1998). Valdemeca (n°9), Riba de Santiuste-Tordelrábano (n°5), Rillo de Gallo (n°6) and Alcázar de San Juán (n°13); Sauropterygia – Sauroptergygia is the best known and by cyamodontids in the Middle Muschelkalk of of marine reptiles of the Iberian Peninsula. It Alpera (n°14) and in the Carnian of Manzarena (n°8). includes remains of Triassic , Finally, indeterminate remains of have been placodonts, , and pistosaurs, and of Jurassic unearthed in the Muschelkalk of Torrijas (n°8), Royuela and Cretaceous plesiosaurs. Most of the fossils are (n°7), Cañete and Boniches (n°10), Bienservida- isolated vertebrae and teeth but complete skeletons, Villarrodrigo (n°15), Mont-ral-Alcover (n°4), and in the notably of nothosaurs, do exist also. Muschelkalk /Keuper? of Alcázar de San Juán (n°13).

Pachypleurosaurid fossils have only been found in fossils are well represented in the Iberian the Ladinian of Mont-ral-Alcover (n°4), and in the Peninsula, mainly by isolated vertebrae and teeth but Muschelkalk of Bienservida-Villarrodrigo (n°15) and also by complete skeletons (Sanz et al ., 1993 in et al et al Aiguafreda (n°2) (Sanz ., 1993 in Quesada ., Quesada et al ., 1998; Rieppel y Hagdorn, 1998; 1998; Rieppel y Hagdorn, 1998; Rieppel, 2000).

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Rieppel, 2000; Quesada y Aguera-González, 2005). The Victoria (n°41), Alvaiázere (n°43), Pintanheira (n°41), nothosaurid cymatosauroides Sanz, 1983, the of Condeixa-a-Nova (n°38), and the from the Ladinian of Mont-ral-Alcover (n°4), is the only Toarcian/Aalenian of Tomar (n°44) (Zbyszwesky y marine reptile named on the basis of material from Moitinho de Almeida, 1952 and Ferreira, 1958 in Spain. The other nothosaurid balsami is the Crespo, 2002). It should be noted that: 1) these species best marine reptile known of the Iberian record. It is are currently considered as synonymous of respectively represented by several complete skeletons of different I. communis and S. megalorhinus ; and 2) these ontogenetic stages of development from the identifications are questioned as most of the specimens Muschelkalk of Estada (n°1) and Mont-ral-Alcover are poorly diagnostical. sp. has been (n°4). Lariosaurus calcagnii and sp. have described in the Pliensbachian of Cadima (n°36), and recently been identified in the Ladinian of Mont-ral- the Pliensbachian/Toarcian? of Murtede (n°36), whereas Alcover (n°4). Nothosaurus sp. has been found in the sp. has been found in the Toarcian of Muschelkalk of Royuela (n°7), Bienservida- Condeixa-a-Nova (n°38) and the Toarcian/Aalenian? of Villarrodrigo (n°15), and tentatively in the Muschelkalk Tomar (n°44). Pending a complete revision of all these of Sóller (n°12) and the Carnian of Manzarena (n°8). specimens, they are provisionnally referred here as Indeterminate nothosaurids are known in the Ichthyosaurus sp. and Stenopterygius sp. Finally, Muschelkalk of Bienservida-Villarrodrigo (n°15) and indeterminate ichthyosaurian remains have been found Olesa (n°3), and the Muschelkalk/Keuper? of Henajeros in the Pliensbachian of Villaviciosa (n°16), Ólvega (n°11), Valdemeca (n°9), Rillo de Gallo (n°6) and Riba (n°20), Cadima (n°36) and Agua de Madeiros (n°40) de Santiuste-Tordelrábano (n°5). (Crespo, 2002; Ruiz-Omeñaca et al ., 2006).

Pistosaur remains have been found in the Ladinian Squamata – Marine squamates consist of of Mont-ral-Alcover (n°4) (Sanz et al ., 1993 in Quesada palaeophid snakes and mosasaurid . The et al ., 1998, Rieppel y Hagdorn, 1998; Rieppel, 2000). pachyophiid Simoliophis rochebrunei (= S. delgadoi of Sauvage 1886) has been described in the Iberian plesiosaurs are exclusivelly represented by Cenomanian of Pendão/Figueira and Cacém (n°53) plesiosauroids. Indeed, pliosauroids are currently (Jonet, 1981 in Crespo, 2002). unknown as the teeth assigned to Polytpychodon from the Aptian of Morella (n°32) belong in fact to a Mosasaurids comprise the highest number of generic baryonychine theropod . sp. has and specific taxa of marine reptiles known in the Iberian been described in the Pliensbachian/Toarcian? Of Peninsula; isolated diagnostical teeth are the most Alhadas (n°37); however, this attribution must be taken frequent remains found. Most remains come from the with caution as the material has not been reviewed since Basque-Cantabrian region. The of Bóveda Sauvage (1897-1898). Cimoliosaurus sp. has been (n°24), Castillo-Lasarte (n°25) and Jáuregui (n°26) have described by Sauvage (1897-1898) in the Cenomanian yielded lemonnieri , Tylosaurus sp., of Alcantara (n°54) but this vertebra rather reminds Leiodon sp. and undeterminated mosasaurines (Bardet those of polycotylids (pers. obs.). Elasmosaurids have et al ., 1997). The Maastrichtian of Laño (n°27) has been found in the Aptian of Morella (n°32) and in the produced Prognathodon solvayi , Leiodon anceps , Maastrichtian of Laño (n°27) (see Bardet et al ., 2008). Platecarpus cf. ictericus , Leiodon sp., Mosasaurus sp. Plesiosauroids have been reported from the Hettangian- and undeterminated mosasaurines (Bardet et al ., 1997). and Pliensbachian of Villaviciosa (n°16), The Campanian-Maastrichtian of Aveiro (n°49) and and from the Kimmeridgian of Colunga (n°17) and Chousa do Fidalgo (n°50) includes Mosasaurus sp. and Ribadesella (n°18) (Ruiz-Omeñaca et al ., 2006). Mosasauridae indet. respectively (Sauvage, 1897-1898, Indeterminate remains of have also been Antunes y Broin, 1988 in Crespo, 2002). Finally, two reported from the Kimmeridgian/Tithonian of Lourinhã vertebrae from the Campanian-Maastrichtian of Viso (n°46), the of Galve (n°29), Castellote and (n°51) described by Sauvage (1897-1898) as Crocodilus Las Parras de Castellote (n°30), the Aptian of Todolella blavieri are probably referable to plioplatecarpine (n° 32), and finally the Mid Cretaceous of São Pedro do mosasaurids (pers. obs.). Estoril (n°53). Crocodyliformes – The only sure occurrence of – Ichthyosauria is the poorest marine crocodiles in the Mesozoic of the Iberian known clade of Iberian marine reptiles. Nearly all Peninsula is that of . Indeed, the presence specimens found are poorly diagnostical remains, i.e. of pholidosaurids in the Hauterivian-Barremian of isolated vertebrae and fragments. The only Triassic Galve (n°29) and the Wealden of Ortigosa de Cameros record is from the Carnian of Manzarena (n°8). All (n°28) needs to be confirmed and that of dyrosaurids in other discoveries are from the Jurassic of both Spain the Cenomanian of Nazaré (Leiria) is doubtful. and Portugal; no Cretaceous record is currently known. Thalattosuchians are relatively well represented in Spain Two species, Ichthyosaurus intermedius and and Portugal by both teleosaurids and metriorhynchids Stenopterygius aff. uniter ., have been repeatedly (Buscalioni, 1986 in Quesada et al ., 1998; Antunes, mentioned in the Lias of the Central region of Portugal, 1967 and Krebs, 1968 in Crespo, 2002; Sauvage, 1897- namely from the Sinemurian of Foz da Ribeira (n°40) 1898). Most specimens consists of isolated vertebrae and Praia N.S. da Victoria (n°41), the Pliensbachian of and jaw fragments, but more complete remains do also Casal Comba (n°35), Murtede (n°36), Praia N.S. da exist.

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The Teleosauridae could be represented by three plesiosauroid, elasmosaurid and ?polycotylid species of Steneosaurus . However, this attribution plesiosaurs), ichthyopterygians (ichthyosaurians), (except S. bollensis ) should be taken with caution squamates (mosasaurids, pachyophiid snakes), because no revision work is known since Sauvage crocodyliforms (teleosaurid and metriorhynchid (1897-1898). They are: cf. S. bollensis in the thalattosuchians; ?pholidosaurids) and turtles Toarcian/Aalenian of Tomar (n°44), S. jugleri in the (plesiochelyid and other chelonioids). Kimmeridgian of Fervença (n°41) and the Tithonian of Praia de Santa Cruz (n°46), and S. cf. obtusidens in the ACKNOWLEDGMENTS Wealden of Benagéber (n°34). Teleosaurids are also known by Teleosaurus sp. from the Jurassic/Cretaceous Financial support by the Ministerio de Educación y of Buñol (n°23), Machimosaurus hugii from the Ciencia (projects CGL2004-02338 and CGL2007- Kimmerigian of Guimarota (n°42) and Machimosaurus 64061/BTE), the Basque Gouvernment (research group sp. from the Kimmeridgian of Colunga (n°17), IT-361-07) and the Principality of Asturias (Protocolo Ribadesella (n°18), Malhão (n°48) and Cesareda (n°45), CN-04-226). This work is a contribution to the the Tithonian of Praia de Santa Cruz (n°46), the “Convenio específico de colaboración” between the Hauterivian-Barremian of Galve (n°29), and the UPV/EHU, CNRS and MNHN. Wealden of Benagéber (Valencia). Undeterminated teleosaurids are known in the Kimmeridgian of Colunga BIBLIOGRAPHY (n°17), Ribadesella (n°18), Cezimbra (n°38), and the Malm of Pombal (n°39). Bardet, N., Corral, J.C. y Pereda Suberbiola, X. (1997): Les Mosasaures (Squamata) du Crétacé supérieur The Metriorhynchidae are represented by remains du Bassin Basco-Cantabrique. Géobios , 20: 19-26. referred to as Metriorhynchus sp. from the Callovian of Bardet, N., Fernández, M., García-Ramos, J.C., Pereda Ricla (n°21) and the Kimmeridgian of Fervença (n°41), Suberbiola, X., Piñuela, L., Ruiz-Omeñaca, J.I. y as well as by ? Dakosaurus from the Wealden of Morella Vincent, P. (2008): A juvenile plesiosaur from the (n°32). Undeterminated thalattosuchians have been Pliensbachian (Lower Jurassic) of Asturias, Spain. mentioned in the Aalenian- of Obón (n°22), the Journal of , 28:258-263 Callovian of Valgañón (n°19), the Kimmeridgian of Crespo, E.G. (2002): Paleoherpetofauna Portuguesa. Colunga (n°17) and Ribadasella (n°18), and the Revista Española de Herpetología , 17-35. Wealden of Alcalá de la Selva (n°31). Quesada, J.M., de la Fuente, M., Ortega, F. y Sanz, J.L. (1998): Bibliografía del registro español de Testudines – Mesozoic marine turtles (Chelonioidea vertebrados mesozoicos. Boletín de la Real cryptodires) are poorly represented in the Iberian Sociedad Española de Historia Natural (Sección Peninsula, especially in Spain, but the specimens are Geología) , 94:101-137. often well preserved, including complete carapaces. Quesada, J.M. y Aguera González, S. (2005): Descripción del primer ejemplar de Ceresiosaurus The Jurassic Plesiochelyidae are the best known and (Reptilia: Sauropterygia), hallado en la Peninsula are represented by Craspedochelys cf. jaccardi in the Iberica en el Ladiniense (Triasico medio) de Mont- Kimmerigian of Romão (n°41) and Craspedochelys ral-Alcover (Tarragona). Estudios Geológicos , 61: choffati in the Tithonian of Vila Franca do Rosário 247-269. (n°47) (Sauvage, 1897-1898; Antunes et al ., 1988 in Rieppel, O. y Hagdorn, H. (1998): Fossil reptiles from Crespo, 2002). Plesiochelyd fossils have recently been the Spanish Muschelkalk (Mont-ral-Alcover, mentioned in the Kimmeridgian of Colunga (n°17) and Province Tarragona). Historical Biology , 13(1): 77- Ribadesella (n°18) (Ruiz-Omeñaca et al ., 2006). 97. Rieppel, O. (2000): Handbuch der Paläoherpetologie. Cretaceous chelonioids are known by a Sauropterygia I: Placodontia, Pachypleurosauria, "Toxochelyidae"-like taxon in the Barremian of Las Nothosauroidea, . Verlag Dr. F. Hoyas (n°33) and by fragmentary remains of cf. Pfeil, München, 134 p. Allopleuron in the Santonian of San Pantaleón de Losa Ruiz-Omeñaca, J.I, García-Ramos, J.C, Piñuela, L., (n°24) (Jimenez-Fuentes, 1995 and Bardet et al ., 1993 Bardet, N., Bermudez-Rochas, D.D., Canudo, J.I. y in Quesada et al ., 1998). Pereda Suberbiola, X. (2006): Restos directos de vertebrados del Jurásico de Asturias. En: CONCLUSION Fernández-Martínez E. (Ed.), XXII Jornadas de la Sociedad Española de Paleontología , 171-172. The fossil record of Mesozoic marine reptiles from Sauvage, H.E. (1897-1898): Les Vertébrés fossiles du the Iberian Peninsula (whis is represented by more than Portugal. Contributions à l'étude des poissons et des 50 sites) extends from the Middle Triassic to the latest reptiles du Jurassique et du Crétacé. Mémoires et Cretaceous; the Middle Triassic (Muschelkalk facies), Communications du Service géologique du the Lower and Upper Jurassic and the Portugal , 1-46. formations being the richnest. Almost all major clades Schulz, G. (1858): Descripción geológica de la are represented: thalattosaurs, sauropterygians provincia de Oviedo . Imprenta y Librería de Don (pachypleurosaurids; placodontoid, placochelyid and José Gonzalez, Madrid, 138 p. cyamodontid placodonts; nothosaurids; pistosaurids;

Geo-Temas 10, 2008 (ISSN: 1567-5172)