A hypertrophied ungual phalanx from the lower Barremian of Spain: Implications for the diversity and palaeoecology of Spinosauridae (Theropoda) in Iberia
Introduction
Megalosauroidea forms a clade of large-bodied theropod dinosaurs whose presence in the Lower Cretaceous of Europe is limited to the specialized clade of Spinosauridae (e.g., Charig and Milner, 1986, Buffetaut, 2007, Canudo et al., 2008, Mateus et al., 2011). Spinosaurids have been described as “crocodile mimics”, being predominantly fish-eating predators with a semi-aquatic lifestyle (Sereno et al., 1998, Rayfield et al., 2007, Amiot et al., 2010, Ibrahim et al., 2014). They were a highly specialized group with a craniodental morphology clearly distinct from other carnivorous dinosaurs (e.g., Sereno et al., 1998, Rayfield et al., 2007, Hendrickx et al., 2016, Vullo et al., 2016). Baryonyx walkeri, first recovered from the Barremian of England, is the best known spinosaurid taxon hitherto and the only known species of Spinosauridae from Europe (Charig and Milner, 1986). The holotype of Baryonyx consists of a partial skeleton including partial skull and postcranium (Charig and Milner, 1986, Charig and Milner, 1997). Along with its crocodile-like skull, Baryonyx also stands out in having a hypertrophied manual ungual of digit I, which is the meaning of its genus name. Such enlarged thumb claw also appears to be present in other spinosaurid taxa such as Suchomimus tenerensis (Sereno et al., 1998) and possibly Spinosaurus aegyptiacus (Ibrahim et al., 2014). Several taxa of large-bodied tetanurans also developed enlarged manual ungual phalanges (see Fig. 1), including the megalosaurid Torvosaurus (Galton and Jensen, 1979), the allosauroid Chilantaisaurus (Benson and Xu, 2008), all megaraptorids (Porfiri et al., 2014) and the tyrannosauroid Dryptosaurus (Brusatte et al., 2011). Noteworthy among more derived theropods is the gigantic size of the forelimbs of Deinocheirus, terminating in powerful yet poorly recurved claws (Osmólska and Roniewicz, 1970, Lautenschlager, 2014).
Remains referred to Baryonyx are also known from the Lower Cretaceous of Portugal and Spain, though they mostly consist of dental material (e.g., Torcida et al., 1997, Fuentes Vidarte et al., 2001, Mateus et al., 2011). The most complete specimen comes from the Barremian Papo Seco Formation of Portugal and includes both cranial and postcranial material (Buffetaut, 2007, Mateus et al., 2011). The Portuguese material was referred to the species Baryonyx walkeri by Mateus et al. (2011) based on the combination of shared dental characters, in spite the fact that some postcranial differences, considered to be the result of intraspecific variation by these authors, occur. In addition to the presence of Baryonyx walkeri in Iberia, several different tooth morphotypes have been reported, implying the presence of more than one spinosaurid taxon for some authors (e.g., Alonso and Canudo, 2016), although the scarcity of associated postcranial material makes testing this hypothesis a challenge.
This paper aims to describe an enlarged theropod manual ungual recently discovered in an outcrop from the Weald facies of Spain. This new specimen is compared in details with those from other theropods. The implications of this new specimen for our understanding of spinosaurid diversity are also discussed.
Section snippets
Geological and palaeontological context
The manual ungual CSC1-4 was recovered from the fossil locality of Caña Seca 1 in the municipality of Gúdar, Teruel province, Spain (Fig. 2). The study area is located in the northeastern part of the Iberian Chain. The latter was developed during the Palaeogene as a result of the Alpine inversion of the Mesozoic Iberian Rift System (e.g., Salas et al., 2001). The Caña Seca 1 fossil site is part of the El Castellar Formation and is palaeogeographically located within the Galve sub-basin, in the
Material and methods
The fossil claw CSC1-4 (Fig. 3, Fig. 4) was collected by surface prospecting during the 2010 fieldwork campaign by the Aragosaurus-IUCA research team of the University of Zaragoza. Other fossil specimens from the same fossil locality were recovered by surface collection or screen washing (microfossils) between 2008 and 2011. Additional items from the El Castellar Formation mentioned in this paper (see Table 1: numbers MOAL-1/1, SM-2/D1, BNA2-7, SUE1-2, VES4-1) were recovered during the same
Systematic palaeontology
Dinosauria Owen 1842
Theropoda Marsh 1881
Megalosauroidea Fitzinger 1843
Spinosauridae Stromer 1915
aff. Baryonyx sp.
Material, locality and horizon. CSC1-4, a left manual ungual claw (digit I?) from the Caña Seca 1 fossil locality, municipality of Gúdar, Teruel province, Spain. El Castellar Formation, lower Barremian (Lower Cretaceous).
Discussion
The identification of the claw CSC1-4 as a theropod manual ungual rather than a pedal ungual is based on several features, such as a proximal articular surface that is dorsoventrally tall and shows a marked median keel, an oval transverse cross-section, and a strong curvature (Agnolin and Chiarelli, 2010).
The asymmetry in the proximal articular surface (i.e., a dorsoventral ridge slightly offset medially) is used to identify CSC1-4 as a left ungual. Given its large size, the attribution of
Conclusion
The record of an isolated large manual claw in the El Castellar Formation yields new evidence on the common presence of spinosaurids within the vertebrate communities of the Barremian of southwestern Europe (i.e., Iberia). It also provides new data on the presence of a second spinosaurid taxon apart from Baryonyx walkeri, reinforcing the information from the tooth record which already pointed to the coexistence of various spinosaurid species in the Barremian of Europe.
This claw is the only
Acknowledgements
This paper forms part of the project CGL2014-53548, subsidized by the Spanish Ministerio de Economía y Competitividad, the European Regional Development Fund, and the Government of Aragón (“Grupos Consolidados” and “Dirección General de Patrimonio Cultural”). JMG and IDM are supported by the Ministerio de Ciencia, Tecnología e Innovación Productiva, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) of Argentina (Postdoctoral Fellowship). MM-A is supported by the Fundação para
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