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Re-Description of the Sauropod Dinosaur Amanzia (“Ornithopsis

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Re-Description of the Sauropod Dinosaur Amanzia (“Ornithopsis Schwarz et al. Swiss J Geosci (2020) 113:2 https://doi.org/10.1186/s00015-020-00355-5 Swiss Journal of Geosciences ORIGINAL PAPER Open Access Re-description of the sauropod dinosaur Amanzia (“Ornithopsis/Cetiosauriscus”) greppini n. gen. and other vertebrate remains from the Kimmeridgian (Late Jurassic) Reuchenette Formation of Moutier, Switzerland Daniela Schwarz1* , Philip D. Mannion2 , Oliver Wings3 and Christian A. Meyer4 Abstract Dinosaur remains were discovered in the 1860’s in the Kimmeridgian (Late Jurassic) Reuchenette Formation of Moutier, northwestern Switzerland. In the 1920’s, these were identifed as a new species of sauropod, Ornithopsis greppini, before being reclassifed as a species of Cetiosauriscus (C. greppini), otherwise known from the type species (C. stewarti) from the late Middle Jurassic (Callovian) of the UK. The syntype of “C. greppini” consists of skeletal elements from all body regions, and at least four individuals of diferent sizes can be distinguished. Here we fully re-describe this material, and re-evaluate its taxonomy and systematic placement. The Moutier locality also yielded a theropod tooth, and fragmen- tary cranial and vertebral remains of a crocodylomorph, also re-described here. “C.” greppini is a small-sized (not more than 10 m long) non-neosauropod eusauropod. Cetiosauriscus stewarti and “C.” greppini difer from each other in: (1) size; (2) the neural spine morphology and diapophyseal laminae of the anterior caudal vertebrae; (3) the length-to-height proportion in the middle caudal vertebrae; (4) the presence or absence of ridges and crests on the middle caudal cen- tra; and (5) the shape and proportions of the coracoid, humerus, and femur. These anatomical diferences, combined with their discrepancy in stratigraphic age, make it unlikely that C. stewarti and “C.” greppini belong to the same genus, as also supported through our phylogenetic analysis. “C.” greppini cannot be assigned to any other contemporaneous sauropod taxon from Europe, but is diagnosed by an autapomorphic rugosity on the posteromedial margin of the humerus, as well as a unique combination of features. As such, we erect the new genus name Amanzia for the Swiss taxon “Ornithopsis” greppini, augmenting the growing diversity of Late Jurassic European sauropods. Our phylogenetic analysis places it outside of Neosauropoda, either as the sister taxon to that clade, or as a member of Turiasauria. Keywords: Dinosauria, Neosauropoda, Late Jurassic, Switzerland, Mesozoic, Europe, Palaeontology 1 Introduction 2017). Although the Late Jurassic record of Swiss dino- Over the last 30 years, Switzerland has seen an impres- saur tracks has steadily grown, the record of dinosaur sive increase in dinosaur research (for reviews see: Meyer body fossils is still rather limited. Up to now, body fossil and Türing 2003; Meyer and Marty 2014; Marty et al. remains of theropods (teeth), stegosaurs, and sauropods are known, but most of these are fragmentary (e.g. Meyer Editorial handling: M.J. Benton. and Lockley 1996; Meyer and Hunt 1998; Meyer and Türing 2003). Among these Swiss body fossils, the fnds *Correspondence: [email protected] 1 Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity documented herein are therefore exceptional. Science, Invalidenstraße 43, 10115 Berlin, Germany Full list of author information is available at the end of the article © The Author(s) 2020. This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creat iveco mmons .org/licen ses/by/4.0/. 2 Page 2 of 48 D. Schwarz et al. In the 1860s, an accumulation of fossil reptilian bones assigned all of the bones to the theropod dinosaur Meg- was discovered in a limestone quarry in the Basse Mon- alosaurus, and erected the species “Megalosaurus meri- tagne, near the city of Moutier, Bern, northwestern Swit- ani” in honour of one of the patrons of the NMB. zerland (Fig. 1). Te frst bones appearing in the quarry In 1920, Berlin vertebrate palaeontologist Janensch had already been privately sold when the Swiss geologist mentioned the theropod tooth from Moutier in his pub- Jean-Baptiste Greppin got notice of the fnd. Greppin rec- lication on the theropod dinosaur Elaphrosaurus from ognized the material to be mostly dinosaurian, and saved Tendaguru, Tanzania, and re-classifed it as Labrosau- the remaining bones for the Natural History Museum in rus (Janensch 1920). In a letter to Freiherr von Huene Basel (Naturhistorisches Museum Basel [NMB]). Subse- (Huene 1922), Janensch pointed out the sauropodian quently, he described and fgured the bones as a faunal nature of the vertebrae fgured by Greppin (1870). Tis component of the “Virgulien” of the Jura Mountains in stirred Huene’s interest and, invited by Hans Georg Ste- the region of Bern (Greppin 1870). Te only mention of hlin, the then director of the NMB, Huene examined and the original site can be found in his monograph where re-described the material from Moutier. Huene assigned he stated (p. 339): , Le squelette entier de ce gigantesque the majority of the bones to the British sauropod genus reptile a été trouvé dans la carrière hypovirgulienne de la Ornithopsis and named a new species “Ornithopsis” grep- basse montagne de Moutier, d’où l’ on a retiré les pierres pini (Huene 1922). Following Janensch (1920), Huene pour la construction du temple. Une grande partie de ce regarded the single theropod tooth as “Labrosaurus squelette bien curieux avec une dent bien conservée se meriani” (Huene 1922, 1926), and identifed four verte- trouve au Musée de Bâle“ [“Te whole skeleton of this brae from the material as belonging to a crocodylomorph gigantic reptile was found in the "Hypovirgulien" of a (Huene 1922) (Table 2). quarry of the Basse Montagne of Moutier, from which Some years later, Huene (1927a) erected the new genus the stones were removed for the construction of the tem- Cetiosauriscus for the partial sauropod skeleton NHMUK ple. Much of this peculiar skeleton with a well-preserved (previously BMNH) R3078 (=“Cetiosaurus leedsi”, Wood- tooth can be found at the Basel Museum”]. On the basis ward 1905). Huene also included “Ornithopsis” greppini in of a theropod tooth found with the material (Tables 1, 2), this new genus, thus re-combining the British and Swiss and in the context of the still poor knowledge about dino- species as Cetiosauriscus leedsi and Cetiosauriscus grep- saurs in the middle of the nineteenth century, Greppin pini, respectively (Huene 1927a, b). Unfortunately, the Basel Zürich Genf Solothurn Bern Rhein Basel Birs Porrentruy bs Delémont Dou Olten Balsthal Moutier Aare Solothurn s hüs Sc Biel 030 km Fig. 1 Geographic position of the original quarry of A. greppini Late Jurassic Swiss sauropod dinosaur Amanzia (“Ornithopsis”) greppini Table 1 Complete list of material that can be assigned to A. greppini Collection no. Material Assignment Previous mention Individual Annotations to type series M.H. 206 Bone fragment No None Not assigned None M.H. 209 Bone fragment No None Not assigned None M.H. 221 (8th) Caudal vertebra No None Individual A None M.H. 235 Caudal vertebra Syntype Huene (1922: p. 82) No assignment possible 17 cm long, 10 high, Bone is missing M.H. 238 Caudal vertebra, mid-caudal region No None Individual C Bone fragment M.H. 357 was glued to this specimen in 2005 M.H. 239 Caudal vertebra, mid-caudal region Syntype Huene (1922: p. 82, fg. 5) Individual C None M.H. 240 Caudal vertebra, mid-caudal region No None Individual C None M.H. 241 Caudal vertebra Syntype Huene (1922: p. 82) No assignment possible 12.5 cm long, 8.5 cm high, Bone is missing M.H. 242 (17th) Caudal vertebra No None Individual A Pathologic, CT study performed in 2007 M.H. 243 Caudal vertebra, mid-caudal region No None Individual D None M.H. 244 (27th) Caudal vertebra No None Individual A None M.H. 245 (28th) Caudal vertebra Syntype Huene (1922: p. 82) Individual A None M.H. 246 (Possibly fourth) metatarsal Syntype Huene (1922: p. 82, fg. 8) Individual A Determined as posterior caudal vertebra by Huene (1922) M.H. 248 (18th) Caudal vertebra No None Individual A Pathologic, CT study performed in 2007 M.H. 249 (20th) Caudal vertebra with remnant of No None Individual A None transverse process M.H. 250 (25th) Caudal vertebra No None Individual A None M.H. 251 Caudal vertebra, mid-caudal region No None Individual C None M.H. 252 Caudal vertebra, mid-caudal region Syntype Huene (1922: p. 82, fg. 6) Individual C None M.H. 253 (26th) Caudal vertebra Syntype Huene (1922: p. 82) Individual A None M.H. 254 (22nd) Caudal vertebra Syntype Huene (1922: p. 82) Individual A None M.H. 255 Caudal vertebra, mid-caudal region No None Individual D None M.H. 256 (24th) Caudal vertebra No None Individual A Pathologic M.H. 257 (19th) Caudal vertebra No None Individual A None M.H. 258 (21st) Caudal vertebra Syntype Huene (1922: p. 82, fg. 7) Individual A Bone fragment M.H. 324 was glued to this specimen in 2005 M.H. 259 Right ulna Syntype Huene (1922: p.
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