Short communicationNew spinosaurid (Theropoda, Megalosauroidea) remains from the Arcillas de Morella Formation (upper Barremian) of Morella, Spain
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
References (59)
- et al.
Dinosaur and other fossil vertebrates from fluvial deposits in the Lower Cretaceous of southern Tunisia
Palaeogeography, Palaeoclimatology, Palaeoecology
(2000) - et al.
A hypertrophied ungual phalanx from the lower Barremian os Spain: implications for the diversity and paleoecology of Spinosauridae (Theropoda) in Iberia
Cretaceous Research
(2018) - et al.
Evolution of the carnivorous dinosaurs during the Cretaceous: the evidence from Patagonia
Cretaceous Research
(2013) - et al.
Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain
Palaeogeography, Palaeoclimatology, Palaeoecology
(2007) - et al.
New theropod remains from the Tiourarén Formation (Middle Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans
Proceedings of the Geologists' Association
(2015) - et al.
New data on spinosaurid dinosaurs from the Early Cretaceous of the Sahara
Comptes Rendus de l'Académie des Sciences à Paris, Sciences de la Terre et des Planètes
(1998) Redescription de Streptospondylus altdorfensis, le dinosaure théropode de Cuvier, du Jurassique de Normandie
Geodiversitas
(2001)The enigmatic theropod dinosaur Erectopus superbus (Sauvage 1882) from the Lower Albian of Louppyle-Château (Meuse, France)
New material of the theropod Ichthyovenator from Ban Kalum type locality (Laos): implications for the synonymy of Spinosaurus and Sigilmassasaurus and the phylogeny of Spinosauridae
Journal of Vertebrate Paleontology, Program and Abstracts
(2014)- et al.
Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy
Palaeontology
(2002)
The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos
Naturwissenschaften
On the spinosaurid theropod teeth from the early Barremian (Early Cretaceous) Blesa Formation (Spain)
Historical Biology
A new contribution to our knowledge of the large-bodied theropods from the Barremian of the Iberian Peninsula: the “Barranco del Hocino” site (Spain)
Journal of Iberian Geology
First spinosaurid dinosaur from Australia and the cosmopolitanism of Cretaceous dinosaur faunas
Biology Letters
A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the United Kingdom and the relationships of Middle Jurassic theropods
Zoological Journal of the Linnean Society
Chronostratigraphy of the Barremian–Early Albian of the Maestrat Basin (E Iberian Peninsula): integrating strontium-isotope stratigraphy and ammonoid biostratigraphy
Newsletters on Stratigraphy
Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri
BYU Geology Studies
Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull
Journal of Vertebrate Paleontology
Phylogeny of Allosauroidea
Journal of Systematic Palaeontology
The spinosaurid dinosaur Baryonyx (Saurischia, Theropoda) in the Early Cretaceous of Portugal
Geological Magazine
Spinosaurid teeth from the Late Jurassic of Tengaduru, Tanzania, with remarks on the evolutionary and biogeographical history of the Spinosauridae
Los restos directos de dinosaurios terópodos (excluyendo aves) en España
Dinosaurios y otros reptiles mesozoicos de España
Primera evidencia de dientes aislados atribuidos a Spinosauridae (Theropoda) en el Aptiano inferior (Cretácico Inferior) de Europa: Formación Arcillas de Morella (Espana)
Ameghiniana
Redescription of the holotype of Dryptosaurus aquilunguis (Dinosauria: Theropoda) from the Upper Cretaceous of New Jersey
Journal of Vertebrate Paleontology
The phylogeny of Tetanurae (Dinosauria: Theropoda)
Journal of Systematic Palaeontology
Baryonyx, a remarkable new theropod dinosaur
Nature
Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey
Bulletin of the Natural History Museum
A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the “middle” Cretaceous of Morocco
PeerJ
Recently rediscovered baryonychine teeth (Dinosauria: Theropoda): new morphologic data, range extension and similarity to Ceratosaurus
Journal of Vertebrate Paleontology
Cited by (10)
First dinosaur remains from Ireland
2021, Proceedings of the Geologists' AssociationCitation Excerpt :This is the basal condition for theropods and is present in coelophysoids (Spielmann et al., 2007), dilophosaurids (Welles, 1984), other averostran-line taxa such as Sarcosaurus (Ezcurra et al., 2020), as well as ceratosaurians (Madsen and Welles, 2000; O’Connor, 2007). Among tetanurans only some megalosauroids are known to possess such a proximally extended fibular crest (Benson, 2010a; Madsen, 1976; Malafaia et al., 2018). Of these megalosauroids only a few develop a crest with a low anteroposteriorly broad mound, as seen in BELUM K12493, including Megalosaurus, Piatnitzkysaurus, Condorraptor, and Magnosaurus (Benson, 2010a; b, Rauhut, 2005).
New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain)
2020, Cretaceous ResearchCitation Excerpt :Faunal interchanges between Africa and Europe have been proposed for the Early Cretaceous (Canudo et al., 2009; Ezcurra and Agnolín, 2012; Dunhill et al., 2016). This is supported by Iberian spinosaurid material that closely resembles that of Gondwanan taxa, e.g. cervical vertebrae of an indeterminate baryonychine (Malafaia et al., 2018, 2019) and crown ornamentation of isolated baryonychine teeth (e.g., Canudo et al., 2008). Indeed, the isolated baryonychine teeth from Spain share fluted labial and lingual surfaces with Suchomimus from Niger.
A new spinosaurid theropod (Dinosauria: Megalosauroidea) from the upper Barremian of Vallibona, Spain: Implications for spinosaurid diversity in the Early Cretaceous of the Iberian Peninsula
2020, Cretaceous ResearchCitation Excerpt :The fossil record of theropod dinosaurs from the Early Cretaceous of the Iberian Peninsula has been interpreted as closely related to other European correlative faunas, particularly those known from the traditional Weald Clay Formation of Surrey, in the United Kingdom (e.g. Ortega et al., 2006). Scarce and mostly fragmentary spinosaurid specimens have been described in Barremian to Aptian levels of Spain and Portugal, which have mostly been related with the English taxon Baryonyx (Vieira and Torres, 1995; Fuentes Vidarte et al., 1999; Buffetaut, 2007; Mateus et al., 2011) or interpreted as belonging to forms closely related with this taxon (Canudo and Ruiz-Omeñaca, 2003; Malafaia et al., 2018). However, a putative relationship with the Gondwanan taxa has also been suggested based on some isolated elements (Canudo et al., 2008; Sánchez-Hernández et al., 2007).
New theropod remains from the late Barremian (Early Cretaceous) of Eastern Iberian Peninsula
2024, Historical BiologySpinosaurus is not an aquatic dinosaur
2022, bioRxiv