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Postcranial material of Nothosaurus marchicus from the Lower Muschelkalk (Anisian) of Winterswijk, The Netherlands, with remarks on swimming styles and taphonomy

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Abstract

The postcranial morphology of Nothosaurus from Winterswijk is described on the basis of three partially preserved skeletons. Because of the association with a diagnostic cranium, two of these can be confidently assigned to N. marchicus. Preserved humeri are recognized as morphotype II, which confirms the relationship between this humeral morphotype and N. marchicus. The occurrence of an additional nothosaur taxon in the Lower Muschelkalk, strong sexual dimorphism, or a broader size range of N. marchicus is evidenced again. The postcranial morphology of Nothosaurus is compared to published and new data on the basal pistosauroid (cf. Cymatosaurus) from the same locality. Numerous shared morphological characters of N. marchicus, Nothosaurus sp., and the basal pistosauroid from Winterswijk hamper assignment of isolated bones, which is presently only possible if a combination of features on multiple bones from a single individual can be assessed. Ontogenetic stage of the described skeletons is discussed as well. Differences between N. marchicus, the basal pistosauroid, and the morphologically similar, but smaller pachypleurosaur A. heterodontus are mainly related to the morphologies of the humerus, ulna, and the clavicle-interclavicle complex, and thereby indicate different modes of locomotion i.e., swimming styles. The pachypleurosaur and the basal pistosauroid were most likely anguilliform swimmers, whereas Nothosaurus also employed its forelimbs during swimming (paraxial swimming). Taphonomical observations on material of the Winterswijk locality indicate the presence of scavengers, although most disarticulated skeletons were transported by water and decayed and disintegrated in shallow water.

Kurzfassung

Die Morphologie des postcranial Skelettes von Nothosaurus wird anhand dreier unvollständig erhaltener Skelette von der Fundstelle Winterswijk beschrieben. Zwei der Skelettreste wurden mit diagnostischem Schädelmaterial gefunden und können so unzweifelhaft N. marchicus zugeordnet werden. Soweit erhalten, können die Humeri dem Nothosaurus-Morphotyp II zugeschrieben werden, was die bereits zuvor gezogene Verbindung dieses Humerus Morphotyps mit diesem Taxon bestätigt. Auch mit dieser Studie wird auf das Vorkommen eines weiteren Nothosaurus Taxons im unteren Muschelkalk, oder einen deutlichen geschlechtlichen Dimorphismus, oder aber auf ein breiteres Größenspektrum für N. marchicus hingewiesen. Postcraniale Elemente von Nothosaurus werden mit der Morphologie des basalen Pistosauroiden (cf. Cymatosaurus) anhand publizierter Daten aber auch anhand von Neufunden aus Winterswijk verglichen. Zahlreichen morphologischen Ähnlichkeiten stehen einige wenige Unterschiede gegenüber was die Zuordnung isolierter Knochen (Einzelfunden) erschwert oder gar unmöglich macht. Nur die Kombination von Merkmalen unterschiedlicher Knochen eines Individuums ermöglicht eine genauere taxonomische Zuordnung. Des weiteren wird das ontogenetische Alter der Skelette diskutiert. Morphologische Unterschiede zwischen Nothosaurus, dem basalen Pistosauroiden und dem morphologisch ebenfalls ähnlichen, obwohl kleineren Pachypleurosaurier A. heterodontus betreffen vor allem den Humerus, die Ulna und den Clavicula-Interclavicula-Komplex, was auf Unterschiede in der Lokomotion, also im Schwimmstil, hindeutet. Der Pachypleurosaurier A. heterodontus und der basale Pistosauroide waren höchstwahrscheinlich anguilliforme Schwimmer, wohingegen Nothosaurus bereits seine Vorderarme zur Fortbewegung nutzte was auf paraxiales Schwimmen deutet. Taphonomische Beobachtungen an den Skeletten von marinen Reptilien aus der Fundstelle Winterswijk beweisen die Anwesenheit von Aasfressern. Die meisten Skelette wurden jedoch im Wasser transportiert und zerfielen dann langsam im Flachwasser, wo sie schließlich in unterschiedlicher Erhaltung eingebettet wurden.

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Acknowledgments

We would like to thank all members of the Muschelkalk Workgroup, especially Henk Oosterink, for their support, and Sibelco Europe MineralsPlus Winterswijk, which operates the Winterswijk Steen-en Kalkgroeve, and its manager Gerard ten Dolle for access to the quarry and the support they provided on their premises. Furthermore we would like to thank the following institutions and individuals for access to the specimens studied in this work: Henk Oosterink, Winterswijk, The Netherlands; Herman Winkelhorst, Aalten, The Netherlands; Jan van den Berg, Schoonhoven, The Netherlands; Wim Berkelder, Winterswijk, The Netherlands; Reinier van Zelst and John de Vos, National Museum of Natural History Naturalis, Leiden, The Netherlands; Daniela Schwarz-Wings, Museum für Naturkunde, Humboldt-Universität, Berlin, Germany. Special thanks go out to the following staff members of the Steinmann Institute, University of Bonn: Olaf Dülfer for his outstanding preparation work and Georg Oleschinski for the excellent photographies.

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Authors and Affiliations

  1. Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stuttgart, Germany

    Nicole Klein

  2. Division of Paleontology, Steinmann Institute, University of Bonn, Nußallee 8, 53115, Bonn, Germany

    Nicole Klein & P. Martin Sander

  3. Department of Zoology and Laboratory of Ornithology, Palacký University, 17. listopadu 50, 771 46, Olomouc, Czech Republic

    Dennis F. A. E. Voeten

  4. Erve Jenneboer 75, 7623 JH, Borne, The Netherlands

    Jos Lankamp

  5. Kerkstraat 52, 7471 AL, Goor, The Netherlands

    Remco Bleeker

  6. Uhlweg 10, 51429, Bergisch Gladbach, Germany

    Oliver J. Sichelschmidt

  7. Beethovenstraat 57, 7075 BD, Etten, The Netherlands

    Marco Liebrand

  8. Miramar Zeemuseum, Vledderweg 25, 8381 AB, Vledder, Drenthe, The Netherlands

    Dennis C. Nieweg

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Correspondence to Nicole Klein.

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We dedicate this paper to Wim Berkelder, Winterswijk, The Netherlands a passionate private collector of Winterswijk fossils.

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Klein, N., Voeten, D.F.A.E., Lankamp, J. et al. Postcranial material of Nothosaurus marchicus from the Lower Muschelkalk (Anisian) of Winterswijk, The Netherlands, with remarks on swimming styles and taphonomy. Paläontol Z 89, 961–981 (2015). https://doi.org/10.1007/s12542-015-0273-5

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