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A New Species of Cricosaurus (Thalattosuchia, Metriorhynchidae) Based Upon a Remarkably Well-Preserved Skeleton from the Upper Jurassic of Germany

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A New Species of Cricosaurus (Thalattosuchia, Metriorhynchidae) Based Upon a Remarkably Well-Preserved Skeleton from the Upper Jurassic of Germany Palaeontologia Electronica palaeo-electronica.org A new species of Cricosaurus (Thalattosuchia, Metriorhynchidae) based upon a remarkably well-preserved skeleton from the Upper Jurassic of Germany Sven Sachs, Mark T. Young, Pascal Abel, and Heinrich Mallison ABSTRACT Herein we describe a new and exceptionally well-preserved skeleton of the metri- orhynchid thalattosuchian Cricosaurus from the upper Kimmeridgian Torleite Formation of Painten in Bavaria (Southern Germany). The specimen is articulated, shows soft-tis- sue preservation, and represents one of the most complete metriorhynchid skeletons known. The exceptional preservation allows us to explore the morphological variation of the tail region in the Metriorhynchidae, a part of the skeleton that has long been neglected. Based on our description and phylogenetic analyses, we name this speci- men Cricosaurus albersdoerferi sp. nov. Our phylogenetic analyses recover a Crico- saurus subclade composed of four species from Southern Germany and one from Argentina. We provide revised diagnoses for the Southern German members of this subclade, revealing the presence of at least four closely-related Cricosaurus species in the upper Kimmeridgian-early Tithonian of Southern Germany. Interestingly, within this subclade there is evidence of rapid change in tail construction and feeding ecology. However, there is no evidence of sympatry between these taxa, and the two species known from the same ammonite subzone are exclusively found in different northern- Tethys lagoons. Most interesting, however, is the variation in the skulls, dorsal neural spines, the tail displacement units, and flukes between these different species. This previously unexplored variation within Metriorhynchidae hints to differences in locomo- tory abilities between different species. Sven Sachs. Naturkunde-Museum Bielefeld, Abteilung Geowissenschaften, Adenauerplatz 2, 33602 Bielefeld, Germany, and Im Hof 9, 51766 Engelskirchen, Germany. [email protected] Mark T. Young. School of GeoSciences, Grant Institute, University of Edinburgh, James Hutton Road, Edinburgh, EH9 3FE, UK. [email protected] Pascal Abel. Senckenberg Centre for Human Evolution and Palaeoenvironment, Eberhard-Karls- Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Germany [email protected]], Heinrich Mallison. Palaeo3D, Dorfstr. 11, 86641 Rain am Lech, Germany and CeNak, Universität http://zoobank.org/2667E3D2-BC97-48C5-89B6-C5AD87C10D82 Sachs, Sven, Young, Mark T., Abel, Pascal, and Mallison, Heinrich. 2021. A new species of Cricosaurus (Thalattosuchia, Metriorhynchidae) based upon a remarkably well-preserved skeleton from the Upper Jurassic of Germany. Palaeontologia Electronica, 24(2):a24. https://doi.org/10.26879/928 palaeo-electronica.org/content/2021/3371-a-new-cricosaurus-species Copyright: June 2021 Paleontological Society. This is an open access article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ SACHS ET AL.: A NEW Cricosaurus SPECIES Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. [email protected] Keywords: Metriorhynchidae; Cricosaurus; Upper Jurassic; Torleite Formation; Germany; new species Submission: 28 September 2018. Acceptance: 18 May 2021. INTRODUCTION while the holotype of C. medius was thought to be lost until recently (see Sachs et al., 2019; Herrera The Upper Jurassic limestones of the Altmühl et al., 2021a). For over 100 years Cricosaurus was valley in Bavaria (Southern Germany) are well considered to be a subjective junior synonym of known for their exclusively well-preserved verte- Geosaurus (first proposed by Fraas, 1902), until brate fossils (e.g., Arratia et al., 2015). They have Young and Andrade (2009) split the two genera. yielded iconic specimens, such as the dinosaurs Cricosaurus is now known as a geographically Archaeopteryx and Compsognathus; numerous widespread taxon, with fossils discovered in Upper pterosaurs, but also aquatic tetrapods including Jurassic strata of Europe, South America, and turtles, pleurosaurs, ichthyosaurs, and thalattosu- North America, as well as Lower Cretaceous strata chian crocodylomorphs (see Arratia et al., 2015 of western Europe (e.g., Frey et al., 2002; Young and references therein). The first thalattosuchian and Andrade, 2009; Herrera et al., 2013a, 2021a, remains described from these units were named 2021b; Sachs et al. 2020). Crocodilus priscus (now Aeolodon priscus, a teleo- Complete skeletons of Cricosaurus, however, sauroid) and Lacerta gigantea (now Geosaurus are rare. The postcranial anatomy is so far only giganteus, a metriorhynchid) by von Sömmerring known for Cricosaurus suevicus Fraas, 1901 (ini- (1814, 1816). They were also the first thalattosu- tially described as Geosaurus suevicus by Fraas, chian taxa to be formally named in scientific publi- 1901) from the upper Kimmeridgian strata of cations (Young et al., 2018). Nusplingen (Baden-Württemberg, SW Germany) Subsequently, a number of thalattosuchian and C. bambergensis Sachs, Young, Abel, and specimens have been reported from localities in Maillison, 2019 from the upper Kimmeridgian strata the Altmühl valley, the majority of which are metrio- of Wattendorf (Bavaria, Southern Germany). The rhynchids (Frey and Tischlinger, 2015). Metrio- postcranial skeleton of C. araucanensis (Gasparini rhynchidae is a clade of pelagic crocodylomorphs and Dellapé, 1976), from the early late Tithonian of that were geographically widespread in the Late Argentina, is known from incomplete and partially Jurassic (e.g., Frey et al., 2002; Buchy et al., 2006; articulated skeletons (Herrera et al., 2009, 2013b, Gasparini et al., 2006; Young et al., 2010; Herrera 2017). Accordingly, the morphological variation of et al., 2013a, 2015). The metriorhynchids of South- the postcranial skeleton is largely unknown. ern Germany are of special importance because of This research describes and names a new their unique preservation in terms of articulation articulated skeleton (BMMS-BK 1-2, Figure 1) that and completeness. In contrast to this the clay pits is one of the most complete and best-preserved of Peterborough UK yielded hundreds of metrio- Cricosaurus fossils on record. The specimen is on rhynchid fossils that are found disarticulated, often display at the Bürgermeister-Müller-Museum in incomplete, and taphonomically deformed (e.g., Solnhofen (Germany) and derives from the Upper Andrews, 1913; Foffa and Young, 2014; Foffa et Jurassic Torleite Formation. It was found in 2002 al., 2018; MTY pers. obs.). Thus, elucidating the by Raimund Albersdörfer in the Rygol quarry in postcranial morphology of Metriorhynchidae, deter- Painten, Bavaria (Southern Germany). BMMS-BK mining how variable it is, and investigating macro- 1-2 has not been described before and yields new evolutionary trends are frustratingly difficult. information on the cranial and postcranial anatomy The genus Cricosaurus was established by of the genus Cricosaurus, revealing a previously Wagner (1858) for material discovered in the Alt- underappreciated diversity in tail morphology mühl valley. The type species, Cricosaurus ele- within Metriorhynchidae. Furthermore, with the gans (Wagner, 1852), is based on a complete but data gleaned from the specimen we revise the dorsoventrally crushed skull. When Wagner (1858) diagnoses for most of the Cricosaurus species of established the genus Cricosaurus, he also the late Kimmeridgian-to-early Tithonian of South- erected two additional species, C. grandis and C. ern Germany. medius. Today, C. grandis is considered to be a species of Geosaurus (Young and Andrade, 2009), 2 PALAEO-ELECTRONICA.ORG FIGURE 1. Cricosaurus albersdoerferi sp. nov. (BMMS-BK 1-2) holotype, late Kimmeridgian of Painten (Southern Ger- many). Skeleton in lateral view. Scale bar equals 200 mm. Institutional abbreviations. BMMS, Bürgermei- (‘Plattensilex’). The formation ranges from the ster-Müller-Museum Solnhofen, Germany; GPIT, upper Aulacostephanus pseudomutabilis to the Palaeontological collections Tübingen, Sencken- upper Hybonoticeras beckeri Tethyan ammonite berg Centre for Human Evolution and Palaeoenv- Zones (Niebuhr and Pürner, 2014). ironment, Eberhard-Karls-Universität Tübingen, The specimen dates to the Hybnoticeras Tübingen, Germany; NAMU, Naturkunde-Museum beckeri Tethyan ammonite Zone and Lithacoceras Bielefeld, Bielefeld, Germany; NKMB, Naturkunde- ulmense Subzone of the Arnstorf Member in the Museum Bamberg, Bamberg, Germany; NHMUK, eastern Franconian Jura. The Arnstorf Member is Natural History Museum, London, UK; SMNS, characterized by the dominating siliceous lime- Staatliches Museum für Naturkunde Stuttgart, Ger- stones (‘Plattensilex’). Lithographic limestones are many; SNSB-BSPG, Staatliche Naturwissenschaft- less abundant (Schweigert, 2007; Niebuhr and liche Sammlungen Bayerns-Bayerische Pürner, 2014). The matrix BMMS-BK 1-2 is embed- Staatssammlung für Paläontologie und Geologie, ded in is a white-coloured thin and fine-laminated Munich, Germany. siliceous limestone. GEOLOGY AND STRATIGRAPHY PALAEOENVIRONMENT BMMS-BK 1-2 was found in the upper Kimme- The rocks of the Arnstorf Member were ridgian Torleite Formation (Figure 2), which forms deposited in the ‘Painten lagoon’ of
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