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GEOGACETA, 53, 2013

First evidence of stegosaurs (Dinosauria: ) in the , , Asturias, N Spain

Primera evidencia de estegosaurios (Dinosauria: Thyreophora) en la Formación Vega, Kimmeridgiense, Asturias

José Ignacio Ruiz-Omeñaca1, Xabier Pereda Suberbiola2, Laura Piñuela1 and José Carlos García-Ramos1

1 Museo del Jurásico de Asturias (MUJA), 33328 Colunga, Spain. [email protected], [email protected], [email protected] 2 Universidad del País Vasco/Euskal Herriko Unibertsitatea, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, 48080 Bilbao, Spain. [email protected]

ABSTRACT RESUMEN

We describe a caudal centrum from an outcrop of the Vega Se describe un centro vertebral caudal de dinosaurio procedente de un Formation (Kimmeridgian) in Colunga (Asturias Principality, Northern Spain). afloramiento de la Formación Vega (Kimmeridgiense) en Colunga (Asturias). It is very similar to the centra of mid-caudal vertebrae of some stegosaurs, Es muy similar a los centros de vértebras caudales medias de algunos este- like and , which are characterized by the presence gosaurios, como Dacentrurus y Stegosaurus, que se caracterizan por la pre- of well developed haemal processes on the posteroventral corner. Because sencia de procesos hemales posteroventrales, pero no es diagnóstica a nivel this character is not diagnostic to the generic level, the vertebral centrum is de género, por lo que se asigna a indet. Constituye la primera assigned to Stegosauria indet. This is the first evidence of stegosaurs in this evidencia de estegosaurios en esta formación geológica. geological formation. Palabras clave: Península Ibérica, Jurásico Superior, vértebra caudal, Key-words: Iberian Peninsula, Late , caudal , Dacentrurus, Dacentrurus, Stegosaurus. Stegosaurus.

Geogaceta, 53 (2013), 37-40. Fecha de recepción: 12 de julio de 2012 ISSN (versión impresa): 0213-683X Fecha de revisión: 25 de octubre de 2012 ISSN (Internet): 2173-6545 Fecha de aceptación: 30 de noviembre de 2012

Introduction garded all the Iberian material previoulsy nian (Sobral Formation) of Lisbon district assigned to Dacentrutrus armatus (Owen (Mateus et al., 2009). Stegosaurian are relatively fre- 1875) as Dacentrurus sp. In Asturias, stegosaur evidences have quent in the of the Iberian Stegosaurus and Miragaia are exclusive been found in Tereñes and Lastres forma- Peninsula.Three genera have been cited up of : tions (Kimmeridgian), which have provided to now: Dacentrurus Lucas 1902, Stegosaurus cf. ungulatus has been des- prints and trackways assigned to the ichno- Stegosaurus Marsh 1877 and Miragaia Ma- cribed in the Kimmeridgian-lower genus (García-Ramos et al., teus, Maidment et Christiansen 2009. (Alcobaça Formation) of Leiria district, Por- 2008; Lockley et al., 2008).A locality in the Dacentrurus has been described in both tugal (Escaso et al., 2007a; later asigned to Lastres Formation (La Escalera, Villaviciosa) Portugal and Spain. Portuguese remains Stegosaurus armatus Marsh 1877 by Maid- contains the only body fossils described up have been recovered since the 1940’s in the ment et al., 2008), and Stegosaurus sp. has to now, assigned to Dacentrurinae indet. districts of Leiria and Lisboa (Galton, 1991; been described in the upper Kimmeridgian- (see Company et al., 2010: Table 1). see also Escaso et al., 2007b, Maidment et lower Tithonian (Lourinhã Group) of Leiria In this paper we describe an isolated al., 2008 and references). and Lisbon districts (Escaso et al., 2008a, vertebral centrum from the Vega Formation. Spanish remains, discovered since 1995 2008b). This is the first evidence of stegosaurs in to now, come from Valencia, Teruel and, Miragaia longicollum Mateus, Maid- that formation, where, hitherto, only thero- with reservations, Burgos provinces (Com- ment et Christiansen 2009, the and pod and sauropod remains have been de- pany et al., 2010, and references therein). only known species of the genus, comes scribed. Maidment et al. (2008; p. 372) have re- from the upper Kimmeridgian-lower Titho-

Copyright© 2013 Sociedad Geológica de España / www.geogaceta.com 37 GEOGACETA, 53, 2013 J.I. Ruiz-Omeñaca, X. Pereda Suberbiola, L. Piñuela and J.C. García-Ramos

Geographical and geological sured over the posterior end).The preserved surface (Fig. 1B: “ch”, 1D). These protuber- location width of the posterior articular face is 87 ances are called “hemal processes” by Gal- mm, but the estimated total width of this ton (1985; p. 220), and are not developed The vertebral centrum described here articular surface would be 104 mm (calcu- on the anteroventral surface. was found in 2003 by one of us (L.P.) in the lated by duplicating the width of the right In ventral view, the haemal processes removed materials of the Vega Formation half measured from the center of that face). are triangular in shape (Fig. 1D).They meas- that crop out 1.7 km north of the town of So, the centrum is slightly wider than high, ure 25 mm antero-posteriorly, 20 mm la- Colunga (Asturias Principality, Northern and higher than long. teromedially and 8 mm dorsoventrally. Spain), when the access road that leads to Both articular surfaces are slightly con- the Museo del Jurásico de Asturias (Juras- cave, so the centrum is amphicoelous. The Discussion sic Museum of Asturias), opened April 2004, excavation on the posterior surface is was in construction. deeper than on the anterior one, which is MUJA-1271 has been identified as a The Vega Formation is a fluviatile unit nearly flat (Fig. 1A, 1C ). mid-caudal centrum by the presence of 1) in the base of the Ribadesella Group, an The form of both articular faces is not facets for chevrons, and 2) a reduced trans- Upper Jurassic sucession made up by four fully observable, but it seems to be sub- verse process. The anterior most caudals formations (Vega, La Ñora,Tereñes and Las- hexagonal to circular. Concentric rings are have no chevron facets and the posterior tres) that crops out in the Gijón-Villaviciosa developed in the external edges of both ar- most ones have no transverse processes Basin, in the northern sector of Asturias. ticular faces, and are more marked on the (Gilmore, 1914; Galton, 1985; Galton and The Vega Formation has been dated as ventral edge. Upchurch, 2004). Kimmeridgian according to ostracods and The lateral side (Fig. 1B) is dorso-ven- Among , the presence of pro- charophytes found in the sections of Playa trally convex and anteroposteriorly concave, ximodorsal projections in the transverse de Vega and Ribadesella (see Schudack and and small nutrient foramina are randomly process is exclusive of stegosaurs. These Schudack, 2002, and references therein), distributed. On the right side there is a structures are always developed in the an- east of Colunga. transverse process (Fig. 1C: “tp”). It is terior half of the transverse process and so placed very high on the centrum, possibly they permit to orientate the centrum. Systematic palaeontology near the base of the pedicels of the neural The most striking feature of this verte- arch, which are not preserved. bra is the presence of well-developed facets Dinosauria Owen 1842 The transverse process is inclined, so its for chevrons (haemal processes) on the pos- Seeley 1888 anterior edge is above the level of the pos- teroventral corner. Stegosauria Marsh 1877 terior one; it forms an angle of about 30º According to Galton and Upchurch with the horizontal. (2004; p. 353), in stegosaurian caudals “the Stegosauria indet. (Fig. 1) The transverse process is anteroposte- proximal chevron facet is usually much riorly elongated and its anterior edge is bro- smaller than the distal chevron facet and Material and provenance ken. It measures, as preserved, 30 mm an- well separated from it”.This feature and the teroposteriorly, 20 mm lateromedially, and general morphology (and size) of the cau- One isolated caudal centrum from Co- 15 mm dorsoventrally. dal vertebra are coherent with its assign- lunga (Asturias Principality, Spain), housed MUJA-1271 shows evidence of a pro- ment to Stegosauria. In ankylosaurs, the in the Museo del Jurásico de Asturias ximodorsal projection on the transverse chevron facets of the caudal vertebrae are (MUJA-1271).Vega Formation, Upper Juras- process (Fig. 1B, 1C; “dorsal process” in commonly less marked, and there is no ev- sic, Kimmerigdian. Galton and Upchurch, 2004). However, this idence of a proximodorsal projection in the part of the centrum is not well preserved transverse process. Description and the size of this projection can not be The Late Jurassic record of ankylosaurs established, but it will be small, as is the in is rather scarce as compared to MUJA-1271 is a partial centrum that case in Dacentrurus (Galton and Upchurch, that of stegosaurs, with only a few disco- preserves the right lateral and ventral sides, 2004; p. 353). veries, including the material of Dracopelta and the half of the proximal and distal ends. The dorsal side is not preserved, so it is Galton 1980 from the Tithonian of Portugal The missing parts of the vertebra (left half of not known if the neural arch was or was not (but no caudal vertebrae are known; see the centrum and, probably, the neural arch) fused to the centrum. In the dorsal most Galton, 1983). In Asturias, stegosaurs have were destroyed by the construction vehicles preserved part of the right lateral side there been described so far in Late Jurassic sites that excavated the access road to the is a shallow elongated depression over the on the basis of both osteological and ich- MUJA. transverse process. nological evidences, whereas ankylosaurs The matrix that surrounded the centrum The ventral side (Fig. 1D) is lateromedi- are currently unknown. had a small crocodylomorph crown tooth (8 ally flat and anteroposteriorly concave, and Haemal processes were already men- mm high) that was removed during the presents facets for the chevrons. On both tioned and figured by Owen (1875) and preparation. ends, right and left facets are differentiated. Marsh (1880) in the first published descrip- The centrum measures 87 mm in an- The posterior facets are much more marked tion of Omosaurus armatus Owen 1875 and teroposterior length (measured ventrally) than the anterior ones and form true protu- of Stegosaurus ungulatus Marsh 1879. and 100 mm in dorsoventral height (mea- berances or tubercles on the posteroventral Owen (1875; p. 55) wrote on this re-

38 Paleontología First evidence of stegosaurs (Dinosauria: Thyreophora) in the Vega Formation, Kimmeridgian, Asturias, N Spain GEOGACETA, 53, 2013

Fig. 1.- MUJA-1271. Mid-caudal centrum of Stegosauria indet. from Colunga (Asturias, N Spain), in anterior (A), right lateral (B), posterior (C) and ventral (D) views. Scale bar = 50 mm. Abbreviations: ch, processes for chevron; tp, transverse process. Fig. 1.- MUJA-1271. Centro caudal medio de Stegosauria indet. procedente de Colunga (Asturias, norte de España), en vistas anterior (A), lateral derecha (B), posterior (C) y ventral (D). La escala son 50 mm. Abreviaturas: ch, procesos para el chevron; tp, proceso transverso. spect that in mid-caudal vertebrae “hypa- cluding Omosaurus lennieri Nopcsa – Miragaia (type species Miragaia longi- pophyses are more distinctly marked.These 1911 from the lower Kimmeridgian collum in the Upper Jurassic of Portu- articular protuberances form a pair at the [Argiles d’Octeville] of Normandy, gal, see above). hind border of the inferior surface of the , see Galton 1985, 1991), Hoffstetter 1957 (type centrum; the articular tracts at the fore bor- – Loricatosaurus Maidment, Norman, species Omosaurus durobrivensis from the der of that surface are barely defined“. Barrett et Upchurch 2008 (type species [] of Cambridgeshire, Marsh (1880; p. 256) noted that “in the Stegosaurus priscus Nopcsa 1911, from U.K.) has been recently regarded as a median caudals ... the faces for chevrons the Callovian [Oxford Clay] of Cam- (Maidment et al., 2008; but are placed on prominent tubercles on the bridgeshire, U.K.; this species was pre- see Buffetaut and Morel, 2009). postero-inferior surface”. viously referred to Lexovisaurus duro- Other historical species of the The record of stegosaurs in the Middle brivensis (Hulke 1887) by Galton, 1985, stegosaurian genus Omosaurus Owen 1875 and Late Jurassic of Europe is made of sev- and Galton and Upchurch, 2004, see (preoccupied by the Omosaurus eral genera: discussion in Maidment et al., 2008), Leidy 1856) are now also regarded as no- – Dacentrurus (type species Omosaurus – Stegosaurus (Stegosaurus cf. ungulatus mina dubia (see Maidment et al., 2008 for armatus from the Kimmeridgian [Lower and Stegosaurus sp. in the Upper Juras- details): Omosaurus hastiger Owen 1877, ] of , U.K.; in- sic of Portugal, see above), and Omosaurus phillipsi Seeley 1892-1893,

Paleontología 39 GEOGACETA, 53, 2013 J.I. Ruiz-Omeñaca, X. Pereda Suberbiola, L. Piñuela and J.C. García-Ramos

Omosaurus leedsi Seeley in von Huene fig. 2L; more views of the same vertebra in Escaso Santos, F. (2005). Primera cita del dino- 1901, and Omosaurus vetustus Huene Escaso Santos, 2005: plate 12, figs. A-E), saurio Stegosaurus (Ornithischia, Thyreo- 1910. but less complete. Escaso Santos (2005; p. phora) del Jurásico Superior europeo.Di- Caudal vertebrae are known in Dacen- 21) identified this vertebra as possibly the ploma de Estudios Avanzados, Universidad Autónoma de Madrid, 35 p. (unpublished). trurus, Stegosaurus and Loricatosaurus but 11th-12th caudal vertebra because of the not in Miragaia. absence of a “dorsal process” on the trans- Escaso, F., Ortega, F., Dantas, P., Malafaia, E., Pi- mentel, N.L., Pereda-Suberbiola, X., Sanz, J.L., Loricatosaurus has an autapomorphy in verse process (Galton and Upchurch, 2004; Kullberg, J.C., Kullberg, M.C., and Barriga, F. their proximal and mid-caudal vertebrae, p. 353). MUJA-1271 preserves a small (2007a). Naturwissenschaften, 94, 367-374. which have “large proximal chevron facet “dorsal process” (see description), so it is Escaso, F., Ortega, F., Dantas, P., Malafaia, E., that unites with the distal one to give a V- possible that it comes from a more anterior Silva, B., and Sanz, J.L. (2007b). In: Cantera shaped centrum” (Galton and Upchurch, position among the mid-caudal series. Paleontológica (O. Cambra-Moo, C. Martínez- 2004; p. 359 as Lexovisaurus; see Maid- Escaso Santos (2005, plate 2, figs. F-J) Pérez, B. Chamero, F. Escaso, S. de Esteban ment et al., 2008; p. 83). In MUJA-1271, also figured another centrum that he iden- Trivigno and J. Marugán-Lobón, Eds.). Dipu- the proximal chevron facets are poorly de- tifies as a mid-caudal [LHNB(CN)-1-08]. tación Provincial de Cuenca, 157-171. veloped so the vertebra do not pertain to However, it has no tranverse processes and Escaso, F., Ortega, F., Dantas, P., Malafaia, E., Loricatosaurus. it is probably a distal caudal. Silva, B., and Sanz, J.L. (2008a). In: Actas III Whether valid or not, the of The observed characters do not permit Congreso Latinoamericano de Paleontología de Vertebrados, 86. Lexovisaurus durobrivensis does not pre- us to distinguish Dacentrurus mid-caudal serve caudal vertebrae, so it is not possible centra from Stegosaurus ones.As both gen- Escaso, F., Ortega, F., Dantas, P., Malafaia, E., Silva, B., and Sanz, J.L. (2008b). In: Libro de to know if they were like those of Lori- era are present in the Kimmeridgian of the resúmenes. XXIV Jornadas de la Sociedad Es- catosaurus priscus. Iberian Peninsula, MUJA-1271 is assigned pañola de Paleontología, 107-108. MUJA-1271 is similar to the figured to Stegosauria indet. Galton, P.M. (1983). Palaeontographica mid-caudal vertebrae of both Dacentrus ar- Abteilung A, 182, 1-25. matus from the U.K. (Owen, 1875: plate 16, Conclusions Galton, P.M. (1985). Journal of Vertebrate Pale- figs. 1-2; Galton, 1985: figs. 2O, 7H-J, 8J-O, ontology, 5, 211-254. 12U-X) and Stegosaurus ungulatus from the A mid-caudal centrum with peculiar Galton, P.M. (1991). Geologica et Palaeontolo- USA (Marsh, 1880: plate 7, figs. 7-8; posteroventral processes for the chevron is gica, 25, 299-327. Gilmore, 1914: fig. 28; Ostrom and McIn- described. It comes from an outcrop of the Galton, P.M. and Upchurch, P.(2004). In: The Di- tosh, 1999: plates 28, 29). Vega Formation (Upper Jurassic, Kimmerid- nosauria, Second Edition (D.B. Weishampel, Compared to Dacentrus armatus, gian) in Colunga (Asturias Principality, P.Dodson and H. Osmólska, Eds.). University MUJA-1271 is, by its dorsally placed trans- North Spain). It has been identified as be- of California Press, 343-362. verse process, more similar to the caudals longing to a stegosaur, and is the first evi- García-Ramos, J.C., Piñuela, L., Ruiz-Omeñaca, posterior to 6th-7th (Galton, 1985: fig. 8M- dence of this group of dinosaurs in the Vega J.I., and Pereda Suberbiola, X. (2008). In: Libro de resúmenes. XXIV Jornadas de la So- O) than to the 6th-7th caudal itself (Galton, Formation. 1985: fig. 8J-L) or even to the more ante- ciedad Española de Paleontología, 33-34. Gilmore, C.W. (1914). Bulletin of the United rior caudals ones that have no chevron Acknowledgements facets (Galton, 1985: fig. 8A-I). States National Museum, 89, 1-136. Lockley, M.G., García-Ramos, J.C., Piñuela, L., In the Iberian Peninsula several caudal This research was supported by the Govern- and Avanzini, M. (2008). Oryctos, 8, 53-70. vertebrae assigned to Dacentrurus have ment of Spain (Ministerio de Economía y Com- been cited and described (see Galton, 1991, petitividad, projects CGL2010-16447/BTE and Maidment, S.C.R., Norman, D.B., Barrett, P.M., Escaso et al., 2008a for Portugal, and Com- CGL2010-18851/BTE) and the Government of and Upchurch, P. (2008). Journal of System- atic Palaeontology, 6, 367-407. pany et al., 2010:Table 1 and references for the Principality of Asturias (Consejería de Edu- Spain) but none of them are mid-caudals, cación, Cultura y Deporte and Universidad de Marsh, O.C. (1880). The American Journal of Sci- Oviedo protocol CN-04-226). The referees Julio ence and Arts, Third Series, 19, 253-259. so a comparison is not possible. Company (Universidad Politécnica de Valencia) A partial skeleton from Casal Novo lo- Mateus, O., Maidment, S.C.R., and Christiansen, and Peter Galton (University of Bridgeport) made N.A. (2009). Proceedings of the Royal Society cality in Batalha (Leiria district, Portugal), valuable comments that improved the paper. B: Biological Sciences, 276, 1815-1821. described by Escaso et al. (2007a) as Ste- Ostrom, J.H. and McIntosh, J.S. (1999). Marsh’s gosaurus cf. ungulatus, preserves a mid- References dinosaurs. The Collections from . caudal vertebra [Laboratório de História Na- Second edition. Yale University Press, xxiv + tural da Batalha, LHNB(CN)-1-02]. It repre- Buffetaut, E. and Morel, N. (2009). Comptes 388 p. sents, up to now, the only stegosaur mid- Rendus Palevol, 8, 545-549. Owen, R. (1875). Palaeontographical Society caudal described in the Iberian Peninsula. Company, J., Pereda Suberbiola, X., and Ruiz- Monographs, 29, 15-94. MUJA-1271 is very similar to the above- Omeñaca, J.I. (2010). Journal of Iberian Geo- Schudack, U. and Schudack, M. (2002). Revista mentioned vertebra (Escaso et al. 2007a: logy, 36, 243-252. Española de Micropaleontología, 34, 1-18.

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