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Edinburgh Research Explorer Metriorhynchid crocodylomorphs from the lower Kimmeridgian of Southern Germany: evidence for a new large-bodied geosaurin lineage in Europe Citation for published version: Abel, P, Sachs, S & Young, M 2020, 'Metriorhynchid crocodylomorphs from the lower Kimmeridgian of Southern Germany: evidence for a new large-bodied geosaurin lineage in Europe', Alcheringa: An Australasian Journal of Palaeontology, vol. 44, no. 2, pp. 312-326. https://doi.org/10.1080/03115518.2019.1701079 Digital Object Identifier (DOI): 10.1080/03115518.2019.1701079 Link: Link to publication record in Edinburgh Research Explorer Document Version: Peer reviewed version Published In: Alcheringa: An Australasian Journal of Palaeontology General rights Copyright for the publications made accessible via the Edinburgh Research Explorer is retained by the author(s) and / or other copyright owners and it is a condition of accessing these publications that users recognise and abide by the legal requirements associated with these rights. 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Oct. 2021 Alcheringa For Peer Review Only Metriorhynchid crocodylomorphs from the lower Kimmeridgian of Southern Germany: evidence for a new large-bodied geosaurin lineage in Europe Journal: Alcheringa Manuscript ID TALC-2019-0003.R3 Manuscript Type: Standard Research Article Date Submitted by the 27-Nov-2019 Author: Complete List of Authors: Abel, Pascal; Senckenberg Centre for Human Evolution and Palaeoenvironment, Fachbereich Geowissenschaften Sachs, Sven; Naturkundemuseum Bielefeld, Geowissenschaften Young, Mark; University of Edinburgh School of GeoSciences, Metriorhynchidae, Geosaurinae, Jurassic, Weißjura Group, Limestone- Keywords: marl-alternations, Geosaurini, Crocodylomorpha URL: http://mc.manuscriptcentral.com/talc E-mail: [email protected] Page 1 of 173 Alcheringa 1 2 3 4 Metriorhynchid crocodylomorphs from the lower Kimmeridgian 5 6 7 of Southern Germany: evidence for a new large-bodied geosaurin 8 9 10 lineage in Europe 11 12 13 14 15 PASCAL ABEL, SVEN SACHS and MARK THOMAS YOUNG 16 17 18 For Peer Review Only 19 Abel, P., Sachs, S., Young, M.T., 2019. Metriorhynchid crocodylomorphs from the lower Kimmeridgian of 20 21 Southern Germany: evidence for a new large-bodied geosaurin lineage in Europe. Alcheringa XXX, X–X. ISSN 22 23 0311-5518 24 25 26 27 Over the last two centuries, numerous exquisitely preserved thalattosuchian crocodylomorph skeletons have been 28 29 found in the Jurassic strata of Southern Germany. While the majority of these specimens occur in Toarcian and 30 31 upper Kimmeridgian–lower Tithonian deposits, thalattosuchian remains are otherwise rare in strata representing 32 33 different stages of the Jurassic. Here, we describe skeletal elements from two large-bodied thalattosuchians 34 35 attributable to the family Metriorhynchidae — these were recovered from lower Kimmeridgian sediments in 36 37 Bavaria and Baden-Württemberg, respectively. These new metriorhynchid fossils are closely comparable in both 38 39 stratigraphic age and dental morphology, and thus may be congeneric. Furthermore, our phylogenetic analysis 40 41 suggests affinity with metriorhynchid remains from France, Switzerland, and the United Kingdom. We interpret 42 these taxa as members of an as yet unnamed geosaurine metriorhynchid lineage (herein termed the “E-clade”) 43 44 from the Kimmeridgian and Tithonian of Europe, which appears to be related to species of Torvoneustes from 45 46 England and Mexico, and Purranisaurus potens from Argentina, collectively contributing to “Subclade T” of the 47 48 tribe Geosaurini. Finally, the metriorhynchid material described herein suggests preservation as a “bloat and 49 50 float” carcass that underwent diagenetic dispersal within a limestone-marl-alternation deposited in an off-shore 51 52 epicontinental marine environment. 53 54 55 56 Pascal Abel [[email protected]], Senckenberg Centre for Human Evolution and Palaeoenvironment, 57 58 Eberhard-Karls-University Tübingen, Sigwartstraße 10, 72076 Tübingen, Germany. Sven Sachs 59 60 [[email protected]], Naturkunde-Museum Bielefeld, Abteilung Geowissenschaften, Adenauerplatz 2, 33602 1 URL: http://mc.manuscriptcentral.com/talc E-mail: [email protected] Alcheringa Page 2 of 173 1 2 3 Bielefeld, Germany. Mark T. Young [[email protected]], School of GeoSciences, Grant Institute, 4 5 University of Edinburgh, James Hutton Road, Edinburgh, EH9 3FE, United Kingdom. 6 7 8 9 Key words: Metriorhynchidae, Geosaurinae, Geosaurini, Crocodylomormpha, Jurassic, Weißjura Group, 10 11 limestone-marl-alternations. 12 13 14 15 THALATTOSUCHIANS were a clade of predominantly marine crocodylomorphs that 16 17 flourished during the Jurassic and Early Cretaceous (e. g. Andrews 1913, Westphal 1962, 18 For Peer Review Only 19 20 Wilberg 2012, Chiarenza et al. 2015, Fanti et al. 2016). Early in their evolution, 21 22 thalattosuchians split into two subclades: the Teleosauroidea and Metriorhynchoidea (Young 23 24 & Andrade 2009). In Germany, teleosauroids are well known from abundant skeletons found 25 26 27 in the Holzmaden area of Baden-Württemberg (e.g. Westphal 1962). Metriorhynchoids are 28 29 comparatively less common, with the most complete remains discovered in the upper Upper 30 31 Jurassic limestones of Southern Germany (e.g. Fraas 1901, 1902, Young & Andrade 2009, 32 33 Sachs et al. 2019a). Fragmentary metriorhynchid skeletons have also been documented from 34 35 36 the Middle Jurassic (e.g. Michelis et al. 1996, Waskow et al. 2017) and Lower Cretaceous 37 38 (Koken 1883, Schröder 1921) of western and northern Germany. 39 40 In Southern Germany, thalattosuchian fossils are mainly restricted to Toarcian and upper 41 42 43 Kimmeridgian/lower Tithonian strata (e.g. Wagner 1852, Fraas 1902, Westphal 1962, 1965, 44 45 Sachs et al. 2019a, b). Dietl & Etzold (1977), Abel et al. (2018), and Aiglstorfer et al. (2019) 46 47 additionally reported fragmentary material from the Aalenian of Baden-Württemberg. 48 49 50 Wellnhofer (1978) described several vertebrae from the middle Oxfordian near Sengenthal in 51 52 Bavaria, which he assigned to Steneosaurus Geoffroy Saint-Hilaire, 1825. The currently best- 53 54 preserved specimen is the holotype of the teleosauroid Machimosaurus buffetauti Young et 55 56 al., 2015a from the lower Kimmeridgian Lacunosamergel Formation near Neuffen in Baden- 57 58 59 Württemberg (Young et al. 2014). 60 2 URL: http://mc.manuscriptcentral.com/talc E-mail: [email protected] Page 3 of 173 Alcheringa 1 2 3 Here, we identify two new unpublished metriorhynchid specimens from the Upper Jurassic 4 5 6 lower Weißjura Group of Southern Germany. The first, SMNS 80149, comprises the anterior 7 8 section of adentary with several in situ teeth. The fossil was found by Alfons Berreth in the 9 10 1950s or 1960s on Braunenberg hill near Aalen-Wasseralfingen in eastern Baden- 11 12 Württemberg. The second specimen, PSHME PH1 incorporates disarticulated postcranial 13 14 15 elements and some associated teeth. It was discovered in 1999 by members of the 16 17 Landesverband für Höhlen- und Karstforschung Bayern (LHK) at the Reichold limestone 18 For Peer Review Only 19 quarry near Drügendorf in northern Bavaria (see Weisel 2000, Abel 2016). Both specimens 20 21 22 derive from the upper Subnebrodites planula Tethyan ammonite Biozone and share the same 23 24 diagnostic dental character states. 25 26 27 28 Institutional abbreviations 29 30 BRSMG, Bristol City Museum & Art Gallery, Bristol, UK; CAMSM, Sedgwick Museum of 31 32 33 Earth Sciences, University of Cambridge, Cambridge, UK; DORCM, Dorset County 34 35 Museum, Dorset, UK; UE, The University of Edinburgh, Edinburgh, UK; FAU, Friedrich- 36 37 Alexander-University Erlangen-Nürnberg, Erlangen, Germany; GLHAM, Hunterian Museum, 38 39 40 University of Glasgow, UK; GPIT, Institut und Museum für Geologie und Paläontologie 41 42 Tübingen, Tübingen, Germany; HME, Heimatmuseum Ebermannstadt; IMPRS-TP, 43 44 International Max-Planck Research School for Translational Psychiatry, Munich, Germany; 45 46 47 LEIUG, Department of Geology, University of Leicester, Leicester, UK; LHK, 48 49 Landesverband für Höhlen- und Karstforschung Bayern e.V., Langquaid, Germany; LMU, 50 51 Ludwig-Maximilians-Universität München, Munich, Germany; NHMUK, Natural History 52 53 Museum, London, UK; NMO, Naturmuseum Olten, Olten, Switzerland; OUMNH, Oxford 54 55 56 University Museum of Natural History, Oxford, UK; PSHME, Paläontologische Sammlung 57 58 Heimatmuseum Ebermannstadt, Ebermannstadt, Germany; SMNS, Staatliches Museum für 59 60 Naturkunde Stuttgart, Stuttgart, Germany. 3 URL: http://mc.manuscriptcentral.com/talc E-mail: [email protected] Alcheringa Page 4 of 173 1 2 3 4 5 6 Geological setting 7 8 9 Both SMNS 80149 and PSHME PH1 were found in carbonate rocks of the lower Weißjura 10 11 Group. This Upper Jurassic marine succession comprises limestones, marlstones, and 12 13 dolostones that form the uppermost strata of the Southern German Franconian and Swabian 14 15 16 Jura (Niebuhr & Pürner 2014; Fig. 1). 17 18 For Peer Review Only 19 SMNS 80149 is partly encased in a micritic limestone matrix (=wakestone