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Los Restos Directos De Dinosaurios Terópodos (Excluyendo Aves) En España

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Los Restos Directos De Dinosaurios Terópodos (Excluyendo Aves) En España Canudo, J. I. y Ruiz-Omeñaca, J. I. 2003. Ciencias de la Tierra. Dinosaurios y otros reptiles mesozoicos de España, 26, 347-373. LOS RESTOS DIRECTOS DE DINOSAURIOS TERÓPODOS (EXCLUYENDO AVES) EN ESPAÑA CANUDO1, J. I. y RUIZ-OMEÑACA1,2 J. I. 1 Departamento de Ciencias de la Tierra (Área de Paleontología) y Museo Paleontológico. Universidad de Zaragoza. 50009 Zaragoza. [email protected] 2 Paleoymás, S. L. L. Nuestra Señora del Salz, 4, local, 50017 Zaragoza. [email protected] RESUMEN La mayoría de los restos fósiles de dinosaurios terópodos de España son dientes aislados y escasos restos postcraneales. La única excepción es el ornitomimosaurio Pelecanimimus polyodon, del Barremiense de Las Hoyas (Cuenca). Hay registro de terópodos en el Jurásico superior (Oxfordiense superior-Tithónico inferior), en el tránsito Jurásico-Cretácico (Tithónico superior- Berriasiense inferior) y en todos los pisos del Cretácico inferior, con excepción del Valanginiense. En el Cretácico superior únicamente hay restos en el Campaniense y Maastrichtiense. La mayor parte de las determinaciones son demasiado generales, lo que impide conocer algunas de las familias que posiblemente estén representadas. Se han reconocido: Neoceratosauria, Baryonychidae, Ornithomimosauria, Dromaeosauridae, además de terópodos indeterminados, y celurosaurios indeterminados (dientes pequeños sin dentículos). La mayoría de los restos son de Maniraptoriformes, siendo especialmente abundantes los dromeosáuridos. Las únicas excepciones son por el momento, el posible Ceratosauria del Jurásico superior de Asturias, los barionícidos del Hauteriviense-Barremiense de Burgos, Teruel y La Rioja, el posible carcharodontosáurido del Aptiense inferior de Morella y el posible abelisáurido del Campaniense de Laño. Además hay algunos terópodos incertae sedis, como los "paronicodóntidos" (entre los que se incluye Euronychodon), y Richardoestesia. Palabras clave. Theropoda, Jurásico, Cretácico, España Most fossil remains of theropod dinosaurs from the Iberian Peninsula are isolated teeth and rare post- cranial bones. The only exception is the ornithomimosaur Pelecanimimus polyodon from the Barremian of Las Hoyas (Cuenca). There are records of theropods in the Upper Jurassic (Upper Oxfordian –Lower Tithonian), the Jurassic-Cretaceous transition (Upper Tithonian-Lower Berriasian) and in all the stages of the Lower Cretaceous, with the exception of the Valanginian. In the Upper Cretaceous they are only found in the Campanian and Maastrichtian. Most of the findings are not diagnostic, which makes it difficult identify all the families that may be represented. The groups include Neoceratosauria, Baryonychidae, Ornithomimosauria, Dromaeosauridae, in addition to unidentified theropods and coelurosaurs (represented by small-unserrated teeth). Most of the remains are Maniraptoriformes, especially dromaeosaurids. The only exceptions at the moment are the possible ceratosaur from the Upper Jurassic (from Asturias), the baryonychids from the Hauterivian-Barremian (Burgos, Teruel and La Rioja), the possible carcharodontosaurid from the Lower Aptian (Morella) and the possible abelisaurid from the Campanian of Laño. There are also some theropods incertae sedis, like the “paronichodontids”, among which we include Euronychodon, and Richardoestesia. Key words. Theropoda, Jurassic, Cretaceous, Spain 1 Canudo, J. I. y Ruiz-Omeñaca, J. I. 2003. Ciencias de la Tierra. Dinosaurios y otros reptiles mesozoicos de España, 26, 347-373. INTRODUCCIÓN Los terópodos son dinosaurios generalmente carnívoros, bípedos, y de tamaño variable. Los dinosaurios más típicos de este grupo poseen los miembros anteriores más pequeños que los posteriores, y las manos presentan adaptaciones para mejorar su capacidad en la caza. El pie tiene tres dedos funcionales, con el dedo I generalmente colocado en la parte posterior y por encima del resto de los dedos, y el dedo V ausente o muy reducido. Tienen tendencia a poseer articulaciones intracraneales y neumaticidad en los huesos. La columna vertebral suele ser horizontal, con excepción del cuello que tiene una característica S. Poseen dientes comprimidos lateromedialmente, curvados hacia atrás y con dentículos en los bordes anterior y posterior. En formas derivadas la morfología de los dientes puede variar, por ejemplo perdiéndose los dentículos. También hay taxones que carecen de dientes, y otros con una morfología dental que indica una dieta omnívora o incluso herbívora. Los terópodos tienen una amplia distribución, ya que se encuentran desde el Triásico hasta el final del Cretácico (Gauthier, 1986; Sereno, 1997; Holtz, 1998; Sanz, 2000). Se han propuesto diferentes filogenias que han permitido avanzar en los últimos años sobre la diferenciación de los grandes clados, que son los usados en este trabajo (véase Holtz, 1998). Taxón Material Yacimiento Formación Edad * Referencia (Megalosaurus sp.) diente Ruedes (A) ? Lastres Kimmeridgiense sup- Egozcue,1873 Theropoda indet. Tithónico inf. ? ?Dromaeosauridae dientes Entre Gijón y Lastres Kimmeridgiense sup- Martínez et al., indet. Ribadesella (A) Tithónico inf. 2000a, 2001 Theropoda indet. diente Entre Gijón y Vega Oxfordiense sup- Martínez et al., Ribadesella (A) Kimmeridgiense inf. 2000a. (Theropoda indet.) vértebra playa de la Vega, Vega Oxfordiense sup- Martínez et al., ?Ceratosauria caudal Ribadesella (A) Kimmeridgiense inf. 2000b Tabla 1. Restos directos de terópodos (excluyendo Avialae) en el Jurásico Superior de España. Abreviaturas: A: Asturias. Referencias en la bibliografía. Entre paréntesis están las determinaciones originales de los taxones que se han modificado en este trabajo. *: las dataciones de las Formaciones son de Schudack y Schudack (2002), y de Suarez-Vega (1974) para el diente de Ruedes. Los restos directos de dinosaurios terópodos en España son relativamente abundantes, pero hasta el momento son fragmentarios, con alguna notable excepción. Los dientes aislados son los más abundantes, ya que los terópodos solían tener más de 50 dientes en sus mandíbulas y además, estos dientes, como 2 Canudo, J. I. y Ruiz-Omeñaca, J. I. 2003. Ciencias de la Tierra. Dinosaurios y otros reptiles mesozoicos de España, 26, 347-373. los de otros dinosaurios, eran continuamente reemplazados durante la vida del animal (Currie et al., 1990). Figura 1. Terópodos del Jurásico Superior de España. A: diente de Theropoda indet. (JRVH-20), Formación Vega (Asturias) en vista labial/lingual. B: diente de Theropoda indet. (JRLH-180), Formación Lastres (Asturias) en vista labial/lingual. C-D: vértebra caudal anterior de Ceratosauria? indet., playa de La Vega (Ribadesella, Asturias), en vistas posterior (C) y lateral izquierda (D). Tomados de Martínez et al., 2000a (A-B) y Martínez et al., 2000b (C-D). El objetivo de este trabajo es poner al día los conocimientos sobre los restos directos de los terópodos españoles (excluyendo Avialae), incluyendo únicamente los datos publicados en revistas científicas. Se han recogido todas las citas y se han situado en el actual contexto cronológico y sistemático. Las tablas se han ordenado por la fecha de publicación. En el caso de restos citados en varios trabajos se ha puesto la referencia más moderna. En las Tablas se ha mantenido la determinación original, pero dado que algunas son manifiestamente obsoletas, se discuten en el texto, proponiéndose en su caso otras determinaciones. Se han publicado restos de la provincias de Álava (Condado de Treviño), Asturias, Burgos, Castellón, Cuenca, Huesca, Lérida, La Rioja y Segovia, en yacimientos del Jurásico superior, Cretácico inferior y el final del Cretácico superior. Para una mejor comprensión, se divide el registro en cinco intervalos: Jurásico superior, tránsito Jurásico-Cretácico, "Neocomiense" (Berriasiense-Barremiese), Aptiense-Albiense, Campaniense y Maastrichtiense. 3 Canudo, J. I. y Ruiz-Omeñaca, J. I. 2003. Ciencias de la Tierra. Dinosaurios y otros reptiles mesozoicos de España, 26, 347-373. Figura 2. Terópodos del tránsito Jurásico-Cretácico de España. A-C: diente de Theropoda indet. (GAOO/CL/14), Cuesta Lonsal (Galve, Teruel), A: vista lingual con detalle de los dentículos mesiales y distales, B: vista mesial, C: sección basal. D-G: fragmento de diente de Coelurosauria indet. (MPZ01/98), Las Cerradicas 2 (Galve, Teruel), D: vista labial/lingual, E: sección basal, F: sección apical, G: reconstrucción hipotética del diente completo. H-K: vértebra caudal anterior de Theropoda indet. (MPZ01/97), Carretera (Galve, Teruel), en vistas lateral izquierda (H), posterior (I), 4 Canudo, J. I. y Ruiz-Omeñaca, J. I. 2003. Ciencias de la Tierra. Dinosaurios y otros reptiles mesozoicos de España, 26, 347-373. ventral (J) y dorsal (K). Modificados de Barco y Ruiz-Omeñaca, 2001a (A-G) y Barco y Ruiz- Omeñaca, 2001b (H-K). TEROPODOS DEL JURÁSICO SUPERIOR (Tabla 1, Figura 1) Los restos de terópodos del Jurásico superior de España se han recuperado en los afloramientos que constituyen la "Costa de los Dinosaurios" en Asturias (Martínez et al., 2000a, 2000b, 2001). Es precisamente en este área donde se encontró el primer resto español de dinosaurio publicado. Schulz (1858) cita un diente aislado del Lías de Ruedes que asigna al tiburón Squalus, y que posteriormente es determinado por Egozcue (1873) como del terópodo Megalosaurus (Sanz, 1984; Sanz et al., 1990). A partir del trabajo de Schulz, la edad del Jurásico detrítico en los alrededores de Lastres se consideró tradicionalmente como Lías, hasta comienzos del siglo XX. Sin embargo los invertebrados de los
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