Abstract
Having provided dozens of articulated, and even more disarticulated skeletons of marine reptiles with varying degrees of completeness, the Middle Triassic fossil deposit of Winterswijk, The Netherlands, might be considered the richest within the entire Germanic Basin concerning associated and partially articulated remains. Aside from these remains, a high number of isolated bones are also found. Among the various marine reptile taxa, the pachypleurosaur Anarosaurus and the nothosaur Nothosaurus are the most common. The skeletons and remains show diverse disarticulation patterns. These patterns range from mostly complete articulated skeletons to clustered and isolated bones. Interestingly, a large number of isolated articulated limbs is found, a pattern which is not known from other marine localities. Although numerous reports have been made on the taxa from Winterswijk, the taphonomy of the locality still remains poorly understood. Here, a series of methods are used to describe the taphonomy of 327 marine reptile specimens from the locality of Winterswijk in a qualitative manner. Furthermore, it is tested whether biofilms such as microbial mats could have had an influence on the preferential preservation of several body regions. Upon investigation, it is concluded that the taphonomy of Winterswijk is very complex and indeed depends on various factors such as: current activity, microbial activity, and anatomical differences between taxa. Finally, a total of 12 recurring patterns were identified within the dataset, which could largely be explained by the aforementioned factors.
Similar content being viewed by others
References
Albers, P. 2005. A Placodontoid jaw fragment from the Lower Muschelkalk of Winterswijk (The Netherlands). PalArch’s Journal of Vertebrate Palaeontology 3: 33–36.
Albers, P. 2011. New Nothosaurus skulls from the Lower Muschelkalk of the western Lower Saxony Basin (Winterswijk, The Netherlands) shed new light on the status of Nothosaurus winterswijkensis. Netherlands Journal of Geosciences 90: 15–22.
Albers, P., and O. Rieppel. 2003. A new species of the sauropterygian genus Nothosaurus from the Lower Muschelkalk of Winterswijk, The Netherlands. Journal of Paleontology 77: 738–744.
Beardmore, S.R., and H. Furrer. 2016a. Preservation of Pachypleurosauridae (Reptilia; Sauropterygia) from the Middle Triassic of Monte San Giorgio, Switzerland. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 280: 221–240.
Beardmore, S.R., and H. Furrer. 2016b. Taphonomic analysis of Saurichthys from two stratigraphic horizons in the Middle Triassic of Monte San Giorgio, Switzerland. Swiss Journal of Geosciences 109: 1–16.
Beardmore, S.R., P.J. Orr, T. Manzocchi, and H. Furrer. 2012a. Float or sink: modelling the taphonomic pathway of marine crocodiles (Mesoeucrocodylia, Thalattosuchia) during the death–burial interval. Palaeobiodiversity and Palaeoenvironments 92: 83–98.
Beardmore, S.R., P.J. Orr, T. Manzocchi, H. Furrer, and C. Johnson. 2012b. Death, decay and disarticulation: modelling the skeletal taphonomy of marine reptiles demonstrated using Serpianosaurus (Reptilia; Sauropterygia). Palaeogeography, Palaeoclimatology, Palaeoecology 337: 1–13.
Bickelmann, C., and P.M. Sander. 2008. A partial skeleton and isolated humeri of Nothosaurus (Reptilia: Eosauropterygia) from Winterswijk, The Netherlands. Journal of Vertebrate Paleontology 28: 326–338.
Carter, D.O., D. Yellowlees, and M. Tibbett. 2007. Cadaver decomposition in terrestrial ecosystems. Naturwissenschaften 94: 12–24.
Cheng, Y., X. Wu, and Q. Ji. 2004. Triassic marine reptiles gave birth to live young. Nature 432: 383–386.
Dames, W. 1890. Anarosaurus pumilio nov. gen. nov. sp. Zeitschrift der Deutschen Geologischen Gesellschaft 42: 74–85.
Delsett, L.L., L.K. Novis, A.J. Roberts, M.J. Koevoets, Ø. Hammer, P.S. Druckenmiller, and J.H. Hurum. 2016. The Slottsmøya marine reptile Lagerstätte: depositional environments, taphonomy and diagenesis. Geological Society of London, Special Publications 434: 165–188.
Diedrich, C.G., and H.W. Oosterink. 2000. Bergings- en documentatietechniek van Rhynchosauroides peabodyi (Faber) - sauriërsporen op de grens Boven-Bontzandsteen/Muschelkalk van Winterswijk. Grondboor & Hamer 54: 125–130.
Dülfer, O., and N. Klein. 2006. Studentische Lehrgrabung im Winterswijker Muschelkalk. Der Präparator 52: 90–96.
During, M.A.D., D.F.A.E. Voeten, A.S. Schulp, and J. Reumer. 2017. A possible Pararcus diepenbroeki vertebra from the Vossenveld Formation (Triassic, Anisian), Winterswijk, The Netherlands. Netherlands Journal of Geosciences 96: 63–68.
Eldijk, T. van, G. Goris, A. Haarhuis, J. Lankamp, H. Winkelhorst, J. Reumer, A. Nel, and T. Wappler. 2017. New fossil insects from the Anisian (Lower to Middle Muschelkalk) from the Central European Basin (Germany and The Netherlands). PalZ 91: 185–194.
Faber, F.J. 1958. Fossiele voetstappen in de Muschelkalk van Winterswijk. Geologie en Mijnbouw 20: 317–321.
Furrer, H. 1995. The Kalkschieferzone (Upper Meride Limestone) near Meride (Canton Ticino, Southern Switzerland) and the evolution of a Middle Triassic intraplatform basin. Eclogae Geologicae Helvetiae 88: 827–852.
Galton, P.M. 2005. Bones of large dinosaurs (Prosauropoda and Stegosauria) from the Rhaetic Bone Bed (Upper Triassic) of Aust Cliff, southwest England. Revue de Paléobiologie 24: 51–74.
Gürich, G. 1884. Ueber einige Saurier des oberschlesischen Muschelkalkes. Zeitschrift der Deutschen geologischen Gesellschaft 36: 125–144.
Hagdorn, H., and O. Rieppel. 1999. Stratigraphy of marine reptiles in the Triassic of Central Europe. Zentralblatt für Geologie und Paläontologie, Teil I 1998: 651–678.
Hagdorn, H., and M. Schulz. 1996. Echinodermen-Konservatlagerstätten im Unteren Muschelkalk Osthessens. 1. Die Bimbacher Seelilienbank von Großenlüder-Bimbach. Geologisches Jahrbuch Hessen 124: 97–122.
Hagdorn, H., and T. Simon. 1993. Rinnenbildungen und Emersion in den Basisschichten des Mittleren Muschelkalkes von Eberstadt (Nordbaden). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 189: 119–145.
Hagdorn, H., and T. Simon. 2010. Vossenveld-Formation. Litholex (Lithostratigraphische Einheiten Deutschlands) ID 45: 1–6. https://litholex.bgr.de/gesamt_ausgabe_neu.php?id=45.
Herngreen, G.F.W., J.H.A. van Konijnenburg-van Cittert, H.W. Oosterink, and R.W.J.M. van der Ham. 2005. Nieuwe geologische, palynologische en paleobotanische gegevens (Muschelkalk, Rhaetien-Lias en Oligoceen) uit de steengroeven van Winterswijk. Grondboor & Hamer 59: 84–97.
Hooijer, D.A. 1959. Records of nothosaurians from the Muschelkalk of Winterswijk, Netherlands. Geologie en Mijnbouw 21: 37–39.
Klein, N. 2009. Skull morphology of Anarosaurus heterodontus (Reptilia: Sauropterygia: Pachypleurosauria) from the Lower Muschelkalk of the Germanic Basin (Winterswijk, The Netherlands). Journal of Vertebrate Paleontology 29: 665–676.
Klein, N. 2010. Long bone histology of Sauropterygia from the Lower Muschelkalk of the Germanic Basin provides unexpected implications for phylogeny. PLoS ONE 5: e11613. https://doi.org/10.1371/journal.pone.0011613.
Klein, N. 2012. Postcranial morphology and growth of the pachypleurosaur Anarosaurus heterodontus (Sauropterygia) from the Lower Muschelkalk of Winterswijk, The Netherlands. Paläontologische Zeitschrift 86: 389–408.
Klein, N., and P. Albers. 2009. A new species of the sauropsid reptile Nothosaurus from the Lower Muschelkalk of the western Germanic Basin, Winterswijk, The Netherlands. Acta Palaeontologica Polonica 54: 589–598.
Klein, N., and T.M. Scheyer. 2014. A new placodont sauropterygian from the Middle Triassic of The Netherlands. Acta Palaeontologica Polonica 59: 887–902.
Klein, N., and O.J. Sichelschmidt. 2014. Remarkable dorsal ribs with distinct uncinate processes from the early Anisian of the Germanic Basin (Winterswijk, The Netherlands). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 271: 307–314.
Klein, N., D.F.A.E. Voeten, A. Haarhuis, and R. Bleeker. 2016. The earliest record of the genus Lariosaurus from the early middle Anisian (Middle Triassic) of the Germanic Basin. Journal of Vertebrate Paleontology 36: e1163712.
Klein, N., D.F.A.E. Voeten, J. Lankamp, R. Bleeker, O.J. Sichelschmidt, M. Liebrand, D.C. Nieweg, and P.M. Sander. 2015. Postcranial material of Nothosaurus marchicus from the Lower Muschelkalk (Anisian) of Winterswijk, The Netherlands, with remarks on swimming styles and taphonomy. Paläontologische Zeitschrift 89: 961–981.
Klompmaker, A.A., and R. Fraaije. 2011. The oldest (Middle Triassic, Anisian) lobsters from The Netherlands: taxonomy, taphonomy, paleoenvironment, and paleoecology. Palaeontologica Electronica 14: 1–16.
Koken, E. 1893. Beiträge zur Kenntnis der Gattung Nothosaurus. Zeitschrift der Deutschen Geologischen Gesellschaft 45: 337–377.
Kowal-Linka, M. 2015. Analysis of marrow cavity fillings as a tool to recognise diverse taphonomic histories of fossil reptile bones: implications for the genesis of the Lower Muschelkalk marine bone-bearing bed (Middle Triassic, Żyglin, S Poland). Palaeogeography, Palaeoclimatology, Palaeoecology 436: 64–76.
Kowal-Linka, M., and A. Bodzioch. 2017. Genesis of the Lower Triassic bonebeds from Gogolin (S Poland): the impact of microbial mats on trapping of vertebrate remains. Palaeogeography, Palaeoclimatology, Palaeoecology 466: 38–58.
Lankamp, J. 2002. Bijtsporen op een sauriërbot uit Winterswijk. Grondboor & Hamer 56: 26–27.
Lankamp, J., and H. Winkelhorst. 1998. Van vondst naar verzameling: vondsten van gewervelde dieren uit de Winterswijkse Muschelkalk. Grondboor & Hamer 52: 17–20.
Martill, D.M. 1993. Soupy substrates: a medium for the exceptional preservation of ichthyosaurs of the Posidonia Shale (Lower Jurassic) of Germany. Kaupia 2: 77–97.
Maxwell, E.E., H. Diependaal, H. Winkelhorst, G. Goris, and N. Klein. 2016. A new species of Saurichthys (Actinopterygii: saurichthyidae) from the Middle Triassic of Winterswijk, The Netherlands. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 280: 119–134.
Mayr, F.X. 1967. Paläobiologie und Stratinomie der Plattenkalke der Altmühlalb. Erlanger Geologische Abhandlungen 67: 1–40.
Meyer, H. von. 1847–1855. Zur Fauna der Vorwelt, zweite Abteilung. Die Saurier des Muschelkalks mit Rücksicht auf die Saurier aus Buntem Sandstein und Keuper. Frankfurt: Heinrich Keller.
Meyer, H. von. 1849. Fossile Fische aus dem Muschelkalk von Jena, Querfurt und Esperstädt. Palaeontographica 1: 195–208.
Neenan, J.M., N. Klein, and T.M. Scheyer. 2013. European origin of placodont marine reptiles and the evolution of crushing dentition in Placodontia. Nature Communications 4: 1621. https://doi.org/10.1038/ncomms2633.
Nopsca, F.V. 1928. Palaeontological notes on reptiles. Geologica Hungarica, Series Palaeontologica 1: 3–84.
O’Keefe, F.R., and L.M. Chiappe. 2011. Viviparity and K-selected life history in a Mesozoic marine plesiosaur (Reptilia, Sauropterygia). Science 333: 870–873.
Oosterink, H.W. 1978a. Arthropoda (Geleedpotigen) uit de Onder-Muschelkalk van Winterswijk. Grondboor en Hamer 32: 2–8.
Oosterink, H.W. 1978b. Enkele restanten van het reptiel Placodus uit de Winterswijkse Muschelkalk. Grondboor en Hamer 32: 175–179.
Oosterink, H.W. 1981. De lamellibranchiaten uit de Winterswijkse Onder-Muschelkalk. Grondboor en Hamer 35: 52–60.
Oosterink, H.W. 1986. Winterswijk, geologie deel II: De Trias-periode (geologie, mineralen en fossielen). Wetenschappelijke Mededelingen van de Koninklijke Nederlandse Natuurhistorische Vereniging 178: 1–120.
Oosterink, H.W. 1987. Een insektevleugel uit de Winterswijkse Muschelkalk. Grondboor en Hamer 41: 148–149.
Oosterink, H.W. 1988. Voetsporen van reptielen en amfibieen uit de Winterswijkse Muschelkalk. Grondboor en Hamer 42: 69–78.
Oosterink, H.W. 1990. Problematische sporen in de Winterswijkse Muschelkalk. Grondboor & Hamer 44: 44–49.
Oosterink, H.W. 2001. Rugvinstekels van Acrodus (kraakbeenvis) uit de Winterswijkse Onder-Muschelkalk. Grondboor & Hamer 55: 16–18.
Oosterink, H.W. 2009. Vondst van een degenkrab (Xiphosura, Limulacea) uit de Onder-Muschelkalk (Trias, Anisien) van Winterswijk. Grondboor & Hamer 63: 125–128.
Oosterink, H.W., W. Berkelder, C. de Jong, J. Lankamp, and H. Winkelhorst. 2003. Sauriërs uit de Onder-Muschelkalk van Winterswijk. Staringia 11: 1–146.
Oosterink, H.W., and G.H. Diepenbroek. 1990. Nieuwe vondsten uit de Winterswijkse Trias. Grondboor & Hamer 44: 150–154.
Oosterink, H.W., and W. Poppe. 1979. Vissen en visresten uit de Onder-Muschelkalk van Winterswijk. Grondboor en Hamer 33: 95–112.
Organ, C.L. 2006. Thoracic epaxial muscles in living archosaurs and ornithopod dinosaurs. The Anatomical Record (Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology) 28: 782–793.
Orr, P.J., L.B. Adler, S.R. Beardmore, H. Furrer, M.E. McNamara, E. Peñalver-Mollá, and R. Redelstorff. 2016. “Stick ‘n’ peel”: explaining unusual patterns of disarticulation and loss of completeness in fossil vertebrates. Palaeogeography, Palaeoclimatology, Palaeoecology 457: 380–388.
Reisdorf, A.G., R. Bux, D. Wyler, M. Benecke, C. Klug, M.W. Maisch, P. Fornaro, and A. Wetzel. 2012. Float, explode or sink: postmortem fate of lung-breathing marine vertebrates. Palaeobiodiversity and Palaeoenvironments 92: 67–81.
Reisdorf, A.G., G.S. Anderson, L.S. Bell, C. Klug, A. Schmid-Röhl, H.J. Röhl, M. Jung, M. Wuttke, M.W. Maisch, M. Benecke, D. Wyler, R. Bux, P. Fornaro, and A. Wetzel. 2014. Reply to “Ichthyosaur embryos outside the mother body: not due to carcass explosion but to carcass implosion” by van Loon (2013). Palaeobiodiversity and Palaeoenvironments 94: 487–494.
Rieppel, O. 1989. A New Pachypleurosaur (Reptilia: Sauropterygia) from the Middle Triassic of Monte San Giorgio, Switzerland. Philosophical Transactions of the Royal Society of London (B: Biological Sciences) 323: 1–73.
Rieppel, O. 2000. Sauropterygia I, Encyclopedia of Paleoherpetology, vol. 12A. Friedrich Pfeil: Verlag Dr.
Rieppel, O. 2001. A new species of Nothosaurus (Reptilia: Sauropterygia) from the upper Muschelkalk (lower Ladinian) of southwestern Germany. Palaeontographica (A: Paläozoologie, Stratigraphie) 263: 137–161.
Rieppel, O., and K. Lin. 1995. Pachypleurosaurs (Reptilia: Sauropterygia) from the Lower Muschelkalk, and a review of the Pachypleurosauroidea. Fieldiana: Geology 32: 1–44.
Salisbury, S.W., E. Frey, D.M. Martill, and M.C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formation of north-eastern Brazil. Palaeontographica (A: Paläozoologie, Stratigraphie) 270: 3–47.
Sander, P.M. 1989. The Pachypleurosaurids (Reptilia: Nothosauria) from the Middle Triassic of Monte San Giorgio (Switzerland) with the description of a new species. Philosophical Transactions of the Royal Society of London (B: Biological Sciences) 325: 561–666.
Sander, P.M., N. Klein, P.C.H. Albers, C. Bickelmann, and H. Winkelhorst. 2014. Postcranial morphology of a basal Pistosauroidea (Sauropterygia) from the Lower Muschelkalk of Winterswijk, The Netherlands. Paläontologische Zeitschrift 88: 55–71.
Sander, P.M., O. Rieppel, and H. Bucher. 1997. A new pistosaurid (Reptilia: sauropterygia) from the Middle Triassic of Nevada and its implications for the origin of the plesiosaurs. Journal of Vertebrate Paleontology 17: 526–533.
Schroeder, H. 1914. Wirbeltiere der Rüdersdorfer Trias. Abhandlungen der Preussischen Geologischen Landesanstalt (Neue Folge) 65: 1–98.
Schulp, A.S., R.W. Bleeker, A. Haarhuis, E. van Spronsen, M.A.D. During, G. Goris, P. Kaskes, Y. Matteman, W. Winkelhorst, and H. Winkelhorst. 2017. A tetrapod swimming traceway from the Triassic of Winterswijk, The Netherlands. Netherlands Journal of Geosciences 96: 273–277.
Stigchel, J. van der. 1942. Langs het spoor der eeuwen, een geologische zwerftocht door Nederland. Amsterdam: Scheltema and Holkema.
Stockar, R. 2010. Facies, depositional environment, and palaeoecology of the Middle Triassic Cassina Beds (Meride Limestone, Monte San Giorgio, Switzerland). Swiss Journal of Geosciences 103: 101–119.
Storrs, G.W. 1991. Anatomy and relationships of Corosaurus alcovensis (Diapsida: Sauropterygia) and the Triassic Alcova Limestone of Wyoming. Bulletin of the Peabody Museum of Natural History 44: 1–151.
Voeten, D.F.A.E., P.M. Sander, and N. Klein. 2014. Skeletal material from larger Eusauropterygia (Reptilia: Eosauropterygia) with nothosaurian and cymatosaurian affinities from the Lower Muschelkalk of Winterswijk, The Netherlands. Paläontologische Zeitschrift 89: 943–960. https://doi.org/10.1007/s12542-014-0250-4.
Weigelt, J. 1927. Rezente Wirbeltierleichen und ihre paläobiologische Bedeutung. Leipzig: Max Weg.
Wild, R., and H.W. Oosterink. 1984. Tanystropheus (Reptilia: Squamata) aus dem Unteren Muschelkalk von Winterswijk, Holland. Grondboor en Hamer 38: 142–148.
Ziegler, P.A. 1990. Geological Atlas of Western and Central Europe, 2nd ed. Den Haag: Shell Internationale Petroleum Maatschappij B.V.
Acknowledgements
First of all, we would like to thank all the members of the Werkgroep Muschelkalk Winterswijk for their manifold support, for access to their private collections, for providing any help and further contacts. Discussion with Dennis Voeten and Melanie During contributed greatly to the understanding of the paleoenvironment. We are grateful to Sibelco Europe MineralsPlus Winterswijk, which operates the Winterwijkse Steengroeve, and provided access to the quarry as well as any logistical support possible. We thank Henk Wim van Dorssen and Dennis Nieweg for their kind hospitality at the Museum TwentseWelle, Enschede, and Miramar Zeemuseum, Vledder, respectively, during several visits to the collections under their care. Furthermore, this study would not have been possible without access to numerous private collections. These are the collections of (in alphabetical order): Wim Berkelder (collection now at TwentseWelle, Enschede), Jan van den Berg, Erik Bleeker, Remco Bleeker, Peter Formanoy, Richard de Haan, Adam Haarhuis, Remco Jalink, Charles de Jong, Harry Kasemir (collection now at Naturalis), Jos Lankamp, Marco Liebrand (collection now at Naturalis), Ben Oostendorp, Henk Oosterink, Cornelis Put, Oliver J. Sichelschmidt, Gerben Spanjaard, Herman Winkelhorst, and Bernd Wolfering. We thank Georg Oleschinski (StIPB) for the brilliant photographs of specimens and Olaf Dülfer (StIPB) for excellent preparation of many specimens. Additionally, we thank Mirjam Cahnbley for proofreading several versions of the manuscript. Last, but not least, we would like to thank the reviewers Eberhard Frey and Heinz Furrer, as well as the editor-in-chief Mike Reich, for their comments on a previous version, and their suggestions that greatly improved the final version of the manuscript.
Author information
Authors and Affiliations
Corresponding author
Additional information
Handling Editors: Hans-Dieter Sues and Walter Joyce.
Electronic supplementary material
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Heijne, J., Klein, N. & Sander, P.M. The uniquely diverse taphonomy of the marine reptile skeletons (Sauropterygia) from the Lower Muschelkalk (Anisian) of Winterswijk, The Netherlands. PalZ 93, 69–92 (2019). https://doi.org/10.1007/s12542-018-0438-0
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12542-018-0438-0