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A Three-Dimensional Skeleton of Goniopholididae from the Late Jurassic of Portugal: Implications for the Crocodylomorpha Bracing System

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A Three-Dimensional Skeleton of Goniopholididae from the Late Jurassic of Portugal: Implications for the Crocodylomorpha Bracing System applyparastyle “fig//caption/p[1]” parastyle “FigCapt” Zoological Journal of the Linnean Society, 2019, XX, 1–28. With 10 figures. Downloaded from https://academic.oup.com/zoolinnean/advance-article-abstract/doi/10.1093/zoolinnean/zlz102/5610606 by Auburn University user on 02 November 2019 A three-dimensional skeleton of Goniopholididae from the Late Jurassic of Portugal: implications for the Crocodylomorpha bracing system E. PUÉRTOLAS-PASCUAL1,2,3*, and O. MATEUS1,2, 1Universidade Nova de Lisboa, Faculdade de Ciências e Tecnologia-GeoBioTec, Monte de Caparica, Portugal 2Museu da Lourinhã, Lourinhã, Portugal 3Aragosaurus-IUCA Research group, Zaragoza, Spain Received 8 March 2019; revised 14 August 2019; accepted for publication 15 August 2019 We here describe an articulated partial skeleton of a small neosuchian crocodylomorph from the Lourinhã Formation (Late Jurassic, Portugal). The skeleton corresponds to the posterior region of the trunk and consists of dorsal, ventral and limb osteoderms, dorsal vertebrae, thoracic ribs and part of the left hindlimb. The paravertebral armour is composed of two rows of paired osteoderms with the lateral margins ventrally deflected and an anterior process for a ‘peg and groove’ articulation. We also compare its dermal armour with that of several Jurassic and Cretaceous neosuchian crocodylomorphs, establishing a detailed description of this type of osteoderms. These features are present in crocodylomorphs with a closed paravertebral armour bracing system. The exceptional 3D conservation of the specimen, and the performance of a micro-CT scan, allowed us to interpret the bracing system of this organism to assess if previous models were accurate. The characters observed in this specimen are congruent with Goniopholididae, a clade of large neosuchians abundant in most semi-aquatic ecosystems from the Jurassic and Early Cretaceous of Laurasia. However, its small size, contrasted with the sizes observed in goniopholidids, left indeterminate whether it could have been a dwarf or juvenile individual. Future histological analyses could shed light on this. ADDITIONAL KEYWORDS: crocodiles – Neosuchia – osteoderms – Lourinhã Formation. INTRODUCTION Although the crocodylomorph record is scarce in the Middle Jurassic, new discoveries in Europe appear to Neosuchia was the dominant clade of Crocodylomorpha indicate that it is during this time that many neosuchian in Europe since the Jurassic. It is a clade of lineages appeared and diversified (Young et al., 2016; mesoeucrocodylian crocodylomorphs that appeared Yi et al., 2017). Prior to that time, the Mesozoic was at the beginning of the Jurassic and, with the extant dominated by non-mesoucrocodylian crocodylomorphs, representatives of the crown group Crocodylia, remain such as protosuchians and sphenosuchians (Langston, until the present day. The oldest member of the clade 1973; Tykoski et al., 2002). is the goniopholidid Calsoyasuchus (Tykoski et al., The European representatives of Neosuchia 2002) from the Kayenta Formation of Arizona, United are mainly composed of marine Tethysuchia States (Sinemurian–Pliensbachian, Lower Jurassic). (Pholidosauridae and Dyrosauridae; e.g. Andrade et al., However, the affinity of Calsoyasuchus within 2011; Fortier et al., 2011) and terrestrial to semi-aquatic Neosuchia has recently been challenged and this taxon Atoposauridae, Bernissartiidae, Goniopholididae (e.g. may be related to Hsisosuchidae instead (Wilberg Brinkmann, 1989; Buscalioni et al., 2008; Schwarz- et al., 2019). Therefore, the appearance of Neosuchia Wings et al., 2009b; Gasca et al., 2012; Puértolas- would have occured later during the Middle Jurassic. Pascual et al., 2015b; Guillaume et al., this volume), and Eusuchia (Puértolas-Pascual et al., 2016), the *Corresponding author. E-mail: [email protected] latter being the crocodyliform lineage that includes © 2019 The Linnean Society of London, Zoological Journal of the Linnean Society, 2019, XX, 1–28 1 2 E. PUÉRTOLAS-PASCUAL and O. MATEUS living crocodylians and their extinct relatives. Another deposits of Romania. Aprosuchus could be closely clade with abundant representatives in the Jurassic related to the putative atoposaurids Theriosuchus, Downloaded from https://academic.oup.com/zoolinnean/advance-article-abstract/doi/10.1093/zoolinnean/zlz102/5610606 by Auburn University user on 02 November 2019 and Lower Cretaceous of Europe is Thalattosuchia, Sabresuchus and Knoetschesuchus (Venczel & a group of exclusively marine crocodylomorphs. Codrea, 2019). In this work, we follow the traditional Nevertheless, there is no complete consensus on the phylogenetic framework that includes Theriosuchus, phylogenetic position of Thalattosuchia within (or Sabresuchus, Knoetschkesuchus and Aprosuchus as outside) Neosuchia, which has been a controversial members of the clade Atoposauridae, since it is, at the issue since the first phylogenetic analyses of moment, the most used in the literature. Crocodylomorpha (e.g. Benton & Clark, 1988; Clark, Goniopholididae is a clade of neosuchian 1994; Pol & Gasparini, 2009; Wilberg, 2015; Wilberg crocodylomorphs with a semi-aquatic lifestyle whose et al., 2019). record ranges from the Early Jurassic (or Middle Within terrestrial ecosystems, the typical Jurassic according to Wilberg et al., 2019) to the crocodylomorph fossil assemblage found in Europe Late Cretaceous, with a predominantly Laurasian from the Middle Jurassic to the Early Cretaceous is distribution. Their remains are abundant in the fossil mainly composed of Atoposauridae, Bernissartiidae record of the Late Jurassic and Early Cretaceous of and Goniopholididae (e.g. Brinkmann, 1989; Buscalioni Europe (e.g. Salisbury et al., 1999; Schwarz-Wings et al., 2008; Schwarz-Wings et al., 2009; Gasca et al., et al., 2009b; Andrade & Hornung, 2011; Salisbury & 2012; Puértolas-Pascual et al., 2015b; Tennant et al., Naish, 2011) and the Iberian Peninsula (e.g. Buscalioni 2016a; Guillaume et al., this volume). & Sanz, 1987; Ortega et al., 1996; Schwarz, 2002; Bernissartiidae is a clade of small durophagous Buscalioni et al., 2013; Puértolas-Pascual, Canudo, neosuchians composed of Bernissartia fagesi Dollo, & Sender, 2015a; Guillaume et al., this volume). The 1883 (Buffetaut, 1975; Buscalioni & Sanz, 1990; Norell clade Goniopholididae has also been subjected to a & Clark, 1990) and Koumpiodontosuchus aprosdokiti strong review during the last several years. The genus Sweetman et al., 2014 from the Barremian of Spain Goniopholis has been used as a wastebasket clade for and England. However, isolated teeth assigned to a variety of species based on fragmentary material this clade have been recovered in several sites from (Andrade et al., 2011). However, new discoveries and the Late Jurassic (Brinkmann, 1989; Guillaume et al., phylogenies show us a much more complex scenario this volume) and Early Cretaceous (Buffetaut & Ford, where several new genera and species have been 1979; Buscalioni et al., 2008; Schwarz-Wings et al., erected (e.g. Andrade et al., 2011; Andrade & Hornung, 2009b; Gasca et al., 2012; Puértolas-Pascual et al., 2011; Salisbury & Naish, 2011; Buscalioni et al., 2013; 2015b) of Europe. Puértolas-Pascual et al., 2015a; Martin et al., 2016; Atoposauridae is a clade of dwarf neosuchian Frederickson et al., 2017; Ristevski et al., 2018). These with a record from the Middle Jurassic to the studies show that Goniopholididae is composed of a Upper Cretaceous (Young et al., 2016). Recently, its variety of basal taxa from Asia and North America and phylogenetic framework has been challenged by two derived European clades, one of which includes Tennant et al. (2016b), who removed Theriosuchus the genus Goniopholis and the other comprised of from Atoposauridae and considered this genus as the genera Anteophthalmosuchus and Hulkepholis paraphyletic. In consequence, they created a new (Puértolas-Pascual et al., 2015a). genus, Sabresuchus, for the taxa Sabresuchus (=Theriosuchus) ibericus (Brinkmann, 1989) and Sabresuchus (=Theriosuchus) sympiestodon (Martin CROCODYLOMORPHS OF PORTUGAL et al., 2010). Subsequently, Schwarz et al. (2017) erected The Jurassic of Portugal is rich in fossil dinosaurs the new genus Knoetschkesuchus for the German (e.g. Antunes & Mateus, 2003; Mateus, 2006), but taxon Knoetschkesuchus langenbergensis Schwarz Crocodylomorpha is one of the most dominant et al., 2017, where they also included the Late Jurassic clades in the fossil assemblages from this period species of Portugal, Knoetschkesuchus (=Theriosuchus) (Mateus et al., 2019; Guillaume et al., this volume). guimarotae (Schwarz & Salisbury, 2005). Therefore, The oldest known crocodylomorph in Portugal is the within this new scenario, the genus Theriosuchus marine thalattosuchian Pelagosaurus tomarensis would be restricted to Theriosuchus pusillus Owen, (Ferreira, 1959) from the latest Toarcian or earliest 1879 and Theriosuchus grandinaris Lauprasert et al., Aalenian (Early or Middle Jurassic) of Tomar 2011. However, a small mandible from the Morrison (Antunes, 1967; Parrilla-Bel et al., 2013). This taxon Formation (USA), originally assigned to a juvenile was later re-assigned to Mystriosaurus cf. bollensis goniopholidid (Foster, 2006), has been recently by Antunes, (1967). Other remains correspond to described as Theriosuchus morrisonensis Foster, 2018. isolated teeth assigned to Machimosaurus sp. from In addition, the Aprosuchus ghirai Venczel & Codrea, the Oxfordian of Cesareda, Cesaredas (west-central 2019 has recently been described
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