Abstract
The origin of the turtle body plan remains one of the great mysteries of reptile evolution. To establish the relationships between turtles of different clades in palaeobiological aspects is highly difficult. Here we describe a review of the oldest Mesozoic turtles of Central Europe (Chelyopsis triunguis; Priscochelys hegnabrunnensis) and their morphological relationship with the toothed turtle (Odontochelys semitestacea) of China. P. hegnabrunnensis is a fossil turtle from the Middle Triassic (lower Ladinian) of Hegnabrunn, Germany. It is known from a single, fragmentary shell plate and is considered a very early turtle, potentially the oldest known stem turtle. The fossil provides insights into the early evolution of turtle shells and their transition from a more primitive state. The habitat preferences of Priscochelys hegnabrunnensis remain uncertain. However, given the informative nature of the material available and, since that it was found mixed with marine and terrestrial faunas, it was probably allochthonous. The other older known turtle record is a trackway from the Bunter Sandstone of Thuringia (Germany) and represents the autochthonous ichno-taxon Chelonipus triunguis. Odontochelys semitestacea was a 220-million-year-old specimen excavated in Triassic deposits in Guizhou, China. Odontochelys only possessed the bottom portion of a turtle‘s armour, the plastron. It did not yet have a solid carapace as most other turtles do. Instead of a solid carapace, Odontochelys possessed broadened ribs like those of modern turtle embryos that still have not started developing the ossified plates of a carapace.
References
- Carr, A. (1955) Handbook of Turtles: The Turtles of the United States, Canada, and Baja California. Ithaca and London: Comstock Pub. Associates a Division of Cornell University Press, 542.
- Carroll, R.L. (1988) Vertebrate Paleontology and Evolution. Freeman and Company, New York, 698 pp.
- Dziomber, L., Joyce, W.G. & Foth, C. (2020) The ecomorphology of the shell of extant turtles and its applications for fossil turtles. PeerJ, e10490. https://doi.org/10.7717/peerj.10490
- Gaffney, E.S. (1990) The comparative osteology of the Triassic turtle Proganochelys. Bulletin of the, American Museum of Natural History, New York, 1–263.
- Hutchison, J.H. & Bramble, D.M. (1981) Homology of the plastral scales of the Kinosternidae and related turtles. Herpetologica, 37, 73–85.
- Joyce, W.G. & Gauthier, J.A. (2004) Palaeoecology of Triassic stem turtles sheds new light on turtle origins. Proceedings of the Royal Society B, 271, 1–5. https://doi.org/10.1098/rspb.2003.2523
- Joyce, W.G. & Karl, H.V. (2006) The world’s oldest fossil turtle: fact versus Fiction. In: Danilov, I.G. & Parham J.F. (Eds), Fossil Turtle Research. Russian J. Herpetol., 13, 104–111.
- Karl, H.V. (2005) The Homology of the Supramarginals in Turtles (Reptilia: Chelonii). Studia geologica Salmanticensia 41, 63–75.
- Karl, H.V. (2012) Fossilium Catalogus I: Animalia pars 149. Testudinomorpha I Index Ichnia et Ova Testudinarum fossilium, Testudinata permotriadica. Backhuys Publishers, Margraf Publishers, Leiden/Weikersheim, 1–20 pp.
- Karl, H.V. (2013) Bericht über Gegenstände, Ziele und Ergebnisse der bisherigen Forschungstätigkeit. Universität Göttingen, 1–9.
- Karl, H.V. (2014) Die fossilen Schildkröten vom Doberg bei Bünde. In: Kaiser, M. & Ebel, R. (Eds), Der Doberg bei Bünde. Eine klassische Fundstelle der Paläontologie, 90–102.
- Karl, H.V. & Tichy, G. (2000) Eine neue Schildkröten- Fährtenspezies aus der Unteren Trias Thüringens. Mauritiana (Altenburg), 17 (3), 421–423.
- Karl, H.V., Safi, A., Coufal, J. & Kutzner, H. (2021) Description and identification of a recently discovered steinkern fragment of a marine turtle from the north Bohemian Saxon Cretaceous Basin, Czech Republic (Central Europe). Journal of Global Ecology and Environment, 12 (4), 36–45.
- Karl, H.V., Safi, A. & Tichy, G. (2024a) Cranial anatomy and holotype reconstruction of the Late Cretaceous turtle, Australobaena chilensis from the Quiriquina Formation, Chile. Mesozoic, 1 (4), 474–482. https://doi.org/10.11646/mesozoic.1.4.5
- Karl, H.V., Safi, A. & Tichy, G. (2024b) Review of fossil records of prehistoric reptiles, their distribution, and paleobiogeographic evolution in Pakistan. Scientific Reports in Life Sciences, 5 (3), 1–23. https://doi.org/10.5281/zenodo.13771832
- Karl, H.V., Safi, A., Tichy, G. & Hanold, W. (2025) Palaeobiological Insights into Mesozoic Turtles from Central Europe and Their Morphological Relationship with the South American Matamata Turtle. Journal of Wildlife and Ecology, 9 (1), 52–68.
- Lepper, J., Rambow, D. & Röhling, H.G. (2013) Lithostratigraphie des Buntsandsteins in Deutschland. In: Deutsche Stratigraphische Kommission (Hrsg.; Koordination und Redaktion: Lepper, J. & Röhling, H.G. für die Subkommission Perm–Trias), Stratigraphie von Deutschland XI. Buntsandstein. Schriftenreihe der Deutschen Gesellschaft für Geowissenschaften, 69, 69–149. https://doi.org/10.1127/sdgg/69/2014/69
- Li, C., Wu, X.C., Rieppel, O., Wang, L.T. & Zhao, L.J. (2008) An ancestral turtle from the Late Triassic of southwestern China. Nature, 456, 497–501. https://doi.org/10.1038/nature07533
- Lichtig, A.J., Lucas, S.G., Klein, H. & Lovelace, D.M. (2017) Triassic turtle tracks and the origin of turtles. Historical Biology, 30 (8), 1112–1122. https://doi.org/10.1080/08912963.2017.1339037
- Lilienstern, H.R.v. (1939) Fährten und Spuren im Chirotheriensandstein von Südthüringen. Fortschritte der Geologie und Palaeontologie, XII, 329–341.
- Lovelace, D.M. & Lovelace, S.D. (2012) Paleoenvironments and pleoecology of a Lower Triassic invertebrate and vertebrate ichnoassemblage from the Red Peak Formation (Chugwater Group), Central Wyoming. Palaios, 27, 636–657. https://doi.org/10.2110/palo.2012.p12-011r
- Lucas, S.G., Heckert A.B. & Hunt A.P. (2000) Probable turtle from the Upper Triassic of East-Central New Mexico. Neues Jahrbuch fur Geologie und Palaontologie, 287–300. https://doi.org/10.1127/njgpm/2000/2000/287
- Młynarski, M. (1976) Testudines. Handbuch der Paläoherpetologie, 7, 130 pp.
- Rieppel, O. (2017) Turtles as Helpful Monsters. Indiana University Press, Bloomington and Indianapolis, 208 pp. https://doi.org/10.2307/j.ctt2005ttv
- Rieppel, O. & Reisz, R.R. (1999) The origin and early evolution of turtles. Annual Review of Ecology, Evolution, and Systematics, 30, 1–22. https://doi.org/10.1146/annurev.ecolsys.30.1.1
- Rögl, F. (1999) Mediterranean and Paratethys: facts and hypotheses of an Oligocene to Miocene paleogeography: short overview. Geologica Carpathica, 50, 330–349. https://doi.org/10.1017/CBO9780511542329.002
- Scheyer, T.M. & Sander, P.M. (2007) Shell bone histology indicates terrestrial palaeoecology of basal turtles. Proceeding of the Royal Society B: Biological Sciences, 274, 1885–1893. https://doi.org/10.1098/rspb.2007.0499
- Scheyer, T.M. (2008) Aging the oldest turtles: the placodont affinities of Priscochelys hegnabrunnensis. Naturwissenschaften, 95, 803–810. https://doi.org/10.1007/s00114-008-0386-8
- Scheyer, T.M. (2009) Conserved bone microstructure in the shells of long-necked and short-necked chelid turtles (Testudinata, Pleurodira). Fossil Record, 12 (1), 47–57. https://doi.org/10.1002/mmng.200800009
- Versluys, J. (1914) Über die Phylogenie des Panzers der Schildkröten und über die Verwandtschaft der Lederschildkröte (Dermochelys coriacea). Palaeontologische Zeitschrift, 1 (1), 321–347. https://doi.org/10.1007/BF03160339
- Völker, H. (1913) Über das Stamm-, Gliedmassen-, und Hautskelett von Dermochelys coriacea L. Zoologische Jahrbucher, 33, 431–552.
- Westphal, F. (1975) Bauprinzipien im Panzer der Placodonten (Reptilia triadica). Paläontologische Zeitschrift, 49 (1/2), 97–125. https://doi.org/10.1007/BF02988070
- Wild, R. (1972) Die Wirbeltierfaunen der fränkischen und südalpinen Mitteltrias (ein Vergleich) / The vertebrate faunae of the Franconian and south-alpine Middle Triassic. Zeitschrift der Deutschen Geologischen Gesellschaft, 123 (1), 229–234. https://doi.org/10.1127/zdgg/123/1972/229
- Xing, L., Lockley, M.G., Klein, H., Romilio, A. & Persons IV, W.S. (2024a) A possible turtle swim track from the late Triassic of the Sichuan Basin, China. Historical Biology, 36 (4), 820–824. https://doi.org/10.1080/08912963.2023.2185885
- Xing, L., Romilio, A., Chunyong, C. & Lockley, M.G. (2024b) Turtle tracks from the middle Jurassic Yaopo formation in Beijing, China. Historical Biology, 36 (10), 2102–2107. https://doi.org/10.1080/08912963.2023.2241064
- Xing, L., Chen, Q., Rozario, A.P., Klein, H., Chou, C., Wang, D. & Qi, Q. (2025) First report of turtle and sauropod track assemblages from the Early Jurassic of Guizhou, China. Paläontologische Zeitschrift, 95–111. https://doi.org/10.1007/s12542-025-00716-1
- Zangerl, R. (1969) The turtle shell. In: Gans, C.. Bellairs, A. d’A & Parsons, T.S. (Eds), Biology of the Reptilia. Academic Press, London, 311–339.