Elsevier

Cretaceous Research

Volume 92, December 2018, Pages 174-183
Cretaceous Research

Short communication
New spinosaurid (Theropoda, Megalosauroidea) remains from the Arcillas de Morella Formation (upper Barremian) of Morella, Spain

https://doi.org/10.1016/j.cretres.2018.08.006Get rights and content

Abstract

The record of spinosaurids currently known in Spain is scarce and represented mainly by isolated teeth and other fragmentary skeletal elements. Most of these previously described specimens have been tentatively attributed to Baryonyx, but there are some differences on dental morphology that may indicate the presence of more than one spinosaurid taxon. A set of postcranial osteological remains of theropod dinosaurs from the upper Barremian Arcillas de Morella Formation of Spain is here described. These elements show a combination of characters compatible with Spinosauridae. The specimens from Morella are interpreted as closely related to other Early and early Late Cretaceous spinosaurids, in particular to Baryonyx walkeri from the Barremian of England, Suchomimus tenerensis from the Aptian of Niger, and Sigilmassasaurus from the Cenomanian of Morocco. However, it was identified a combination of features distinct from other spinosaurids, supporting the presence of a distinct form, adding to the knowledge of the diversity and distribution of this clade of theropod dinosaurs in the Lower Cretaceous of the Iberian Peninsula.

Introduction

The Baryonychinae (sensu Sereno et al., 1998) is a group of megalosauroid theropods, which together with Spinosaurinae constitute the clade Spinosauridae. Some recent phylogenetic analyses recover a large polytomy of spinosaurids, casting doubt on the traditional distinction between Baryonychinae and Spinosaurinae (Evers et al., 2015, Sales and Schultz, 2017). Traditionally, the fossil record of baryonychine theropods includes the species Baryonyx walkeri from the lower Barremian of Surrey in England (Charig and Milner, 1986), Suchomimus tenerensis from the Aptian of Tenere Desert in Niger (Sereno et al., 1998), and Ichthyovenator laosensis from the Aptian of Savannakhet Basin in Laos (Allain et al., 2012). The latter was more recently reinterpreted and considered more closely related to Spinosaurinae (Allain, 2014, Hone and Holtz, 2017). In fact, this taxon shows a combination of features shared with traditional baryonychine taxa, together with characters that have been considered typical of spinosaurines (Evers et al., 2015).

Other enigmatic taxon, Sigilmassasaurus brevicollis from the Cenomanian of Morocco, has been more recently considered as a spinosaurid in a mostly consensual way, but with somewhat different taxonomic and systematic interpretations (McFeeters et al., 2013, Ibrahim et al., 2014, Evers et al., 2015). It was proposed as a junior synonym of Spinosaurus aegyptiacus from the Cenomanian of Egypt by Ibrahim et al. (2014), but this synonymy is refuted by Evers et al. (2015). Besides, the description of additional material suggests the presence of more than one spinosaurid taxon in the Kem Kem beds (Evers et al., 2015, Hendrickx et al., 2016).

Some specimens from the upper Apian of Gadoufaoua in Niger were first interpreted as a new spinosaurid, Cristatusaurus lapparenti (Taquet and Russell, 1998). However, other works attributed these specimens to an indeterminate species of Baryonyx (Charig and Milner, 1997) or consider C. lapparenti as a nomen dubium (Sereno et al., 1998).

Beside the mentioned taxa, several isolated remains, mainly teeth, have also been tentatively assigned to this group of specialized theropods. The currently known fossil record of spinosaurids spans from the Barremian of England (Baryonyx walkeri: Charig and Milner, 1997) to the Cenomanian of Egypt (Spinosaurus aegyptiacus: Stromer, 1915) and Brazil (Oxalaia quilombensis: Kellner et al., 2011). However, several putative spinosaurid occurrences, based mainly on isolated teeth, have been described, suggesting a much wider stratigraphic and geographic distribution of the clade (e.g. Benton et al., 2000, Medeiros, 2006, Barrett et al., 2011). Fragmentary remains of possible spinosaurids are known as old as the Middle Jurassic of Niger (Serrano-Martínez et al., 2015) and have been also tentatively identified in the Late Jurassic of the Tendaguru (Buffetaut, 2008; see Rauhut, 2011 for a different interpretation) and Morrison (Allain et al., 2012) formations.

In the Iberian Peninsula, some specimens found in Barremian to Aptian levels of Spain and Portugal, have been referred to baryonychine theropods, some of them tentatively related with Baryonyx walkeri (Vieira and Torres, 1995, Mateus et al., 2011). The Spanish record includes both cranial (mainly isolated teeth) and postcranial material. A fragment of a maxilla found in Barremian – upper Aptian strata of Igea (La Rioja Province) was referred to Baryonyx walkeri (Vieira and Torres, 1995). In addition, an unpublished almost complete hindlimb collected in the same area has also baryonychine affinities and may belong to the same taxon as the maxilla (pers. obs.). In the Cameros Basin, near the locality of Salas de los Infantes (Burgos Province), a set of cranial (a tooth fragment, a postorbital and a squamosal), axial (one cervical, fragments of two sacral, and a caudal vertebrae, some ribs and chevrons), and appendicular (metacarpals and a fragment of a phalanx) elements recovered from upper Hauterivian or lower Barremian deposits were referred to Baryonyx (Fuentes Vidarte et al., 1999). Also, a specimen represented by cervical, dorsal, caudal and sacral vertebrae, the ilia and the ischia of an indeterminate spinosaurid have been described from the Arcillas de Morella Formation at Vallibona, Castellón (Gómez-Fernández et al., 2007). Other isolated remains tentatively attributed to this taxon known in the Spanish fossil record include a manual ungual from El Castellar Formation (lower Barremian) in the Maestrazgo Basin (Gasca et al., 2018) and several teeth from different Lower Cretaceous (late Hauterivian–early Aptian) sites in Teruel, Burgos and Castellón (Torcida et al., 1997, Torcida et al., 2003, Ruiz-Omeñaca et al., 1997, Ruiz-Omeñaca et al., 2005, Canudo and Ruiz-Omeñaca, 2003, Gasulla et al., 2006, Ortega et al., 2006, Sánchez-Hernández et al., 2007, Canudo et al., 2008, Alonso and Canudo, 2016, Alonso et al., 2018). Some of these teeth have been assigned as cf. Baryonyx sp. (Canudo and Ruiz-Omeñaca, 2003). Other teeth, which differ from those of the English species in having distinct ornamented lingual and labial surfaces of the crown and serrations entirely restricted to the distal margin, have been attributed to indeterminate Baryonychinae (Ruiz-Omeñaca et al., 2005, Ortega et al., 2006, Canudo et al., 2008, Alonso and Canudo, 2016). Also some teeth collected in strata of the Camarillas Formation (lower Barremian) in Galve (Teruel Province) were tentatively assigned to spinosaurines based on the lack of serrated carinae and the conical shape of the crowns (Sánchez-Hernández et al., 2007). The identification of different tooth morphotypes may indicate the presence of more than one spinosaurid taxon, but this hypothesis is difficult to test due to the fragmentary nature of the specimens and the lack of detailed descriptions for most of them. On the other hand, some studies of baryonychine teeth from the collections of the Natural History Museum in London (UK) indicate a higher morphological variability on the dentition of these theropods than previously thought (Fowler, 2007).

In Portugal, a set of cranial elements (jaw fragments with teeth) from the Barremian Papo Seco Formation (Cabo Espichel, Setubal), first identified as a new species of Crocodyliformes Suchosaurus girardi Sauvage 1897–98 was later referred to Baryonyx (Buffetaut, 2007). Another specimen collected in Praia dos Lagosteiros (Cabo Espichel, Papo Seco Formation), which includes a partial dentary with teeth, an almost complete tooth with root, some pre-sacral and caudal vertebrae, ribs, pelvic and appendicular elements was referred to Baryonyx walkeri (Mateus et al., 2011).

Isolated post-cranial remains collected in the Mas de la Parreta Quarry, located close to Morella (Castellón Province, East of Spain) are here described. These specimens show morphologies compatible with baryonychine spinosaurids, previously identified in the Iberian record based on both cranial and postcranial elements, and are here tentatively assigned to this clade.

Institutional abbreviations: CMP, Mas de la Parreta Quarry, Morella, Spain; NHMUK, The Natural History Museum, London, U.K.

Section snippets

Materials and methods

The specimens were discovered during paleontological control of the Mas de la Parreta Quarry at Morella (Spain) under permits obtained from the Dirección General de Cultura y Patrimonio [Consellería de Educación, Investigación, Cultura y Deporte-Generalitat Valenciana].

The material here described comprises a mid-cervical vertebra, an almost complete right femur, a proximal fragment of a right femur, and an almost complete left tibia, and are housed at the Museo de la Valltorta (Tirig,

Systematic paleontology

  • Dinosauria Owen, 1842

  • Theropoda Marsh, 1881

  • Tetanurae Gauthier, 1986

  • Megalosauroidea Huxley, 1889

  • Spinosauridae Stromer, 1915

  • Spinosauridae indet.

  • (Fig. 2, Fig. 3, Fig. 4)

  • Referred specimens. CMP-3b/42, a mid-cervical vertebra; CMP-3b/211, an almost complete right femur; CMP-MS-0/22, a proximal fragment of a right femur; CMP-3c/188, an almost complete left tibia.

  • Locality and horizon. Specimens were collected in three different sites (CMP-3b, CMP-3c, and CMP-MS-0) within the Mas de la Parreta Quarry at

Cervical vertebra (CMP-3b/42)

The cervical vertebra, CMP-3b/42 (Fig. 2), is almost complete and well-preserved despite some distortion due to lateral compression, especially on the medial part of the centrum and the posterior articular facet. The centrum is opisthocoelous, with the anterior articular facet strongly convex and the posterior one strongly concave. The anterior articular facet is delimited by a narrow flattened band and has a rounded contour, slightly wider than high (Table 1). The anterior facet is in a dorsal

Discussion

The vertebra, CMP-3b/42, has a general morphology similar to the sixth cervical vertebra of Baryonyx walkeri (Charig and Milner, 1997, Fig. 20E; see Evers et al., 2015 for a re-interpretation of the presacral series of Baryonyx). It shares with most basal tetanurans the presence of a single anterior pleurocoel, contrasting with the two openings generally present in carcharodontosaurids (Carrano et al., 2012). The morphology of the neural spine, thin and with a homogenous anteroposterior length

Conclusions

The set of osteological remains herein described has a combination of morphological characters compatible with Spinosauridae. The cervical vertebra, CMP-3b/42, presents a general morphology similar to the sixth cervical vertebrae of Baryonyx walkeri and a combination of features shared with this taxon and with other spinosaurids, including the elongated zygapophyses, the low and transversely thin neural spine projecting slightly to the rear; and the well-developed epipophyses. However, the

Acknowledgments

We sincerely thank Victor Beltrán, María José Adelantado and Vega del Moll S.A. Company for their involvement and collaboration in the localization of the different fossil sites at the Mas de la Parreta Quarry. We also thank to two anonymous reviewers and to the editors for comments and suggestions to the manuscript. A particular thanks to Serjoscha Evers for his comments and suggestions to the manuscript and also for sharing photos of spinosaurid specimens, which really facilitated a

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