Abstract
We provide a detailed description of the poorly known reptile Eifelosaurus triadicus Jaekel, 1904 from the Upper Buntsandstein (Triassic: early Anisian) of Oberbettingen in the southwestern Eifel region of Rhineland-Palatinate, Germany. The holotype and only known specimen is a partial postcranial skeleton exposed in ventral view. Since its original description Eifelosaurus triadicus has been almost completely ignored in the literature. Our phylogenetic analysis recovered this taxon within Rhynchosauria and Rhynchosauridae, as the sister taxon to the clade comprising Stenaulorhynchinae and hyperodapedontine-line rhynchosaurids. A unique combination of character states allows distinguishing Eifelosaurus triadicus from other rhynchosaur species with preserved matching postcranial bones. Thus, we tentatively consider this taxon valid until more information about the postcrania of other Early to Middle Triassic rhynchosaurs becomes available. Eifelosaurus triadicus represents a paleobiogeographically important record, showing that rhynchosaurs had already attained a broad paleolatitudinal distribution by the early Middle Triassic.
Similar content being viewed by others
References
Abel, O. 1919. Die Stämme der Wirbeltiere. Berlin and Leipzig: Vereinigung wissenschaftlicher Verleger Walter de Gruyter & Co.
Benton, M.J. 1983. The Triassic reptile Hyperodapedon from Elgin: functional morphology and relationships. Philosophical Transactions of the Royal Society of London (B) 302: 605–717.
Benton, M.J. 1985. Classification and phylogeny of the diapsid reptiles. Zoological Journal of the Linnean Society 84: 97–164.
Benton, M.J. 1990. The species of Rhynchosaurus, a rhynchosaur (Reptilia, Diapsida) from the Middle Triassic of England. Philosophical Transactions of the Royal Society of London (B) 328: 213–306.
Bourquin, S., S. Péron, and M. Durand. 2006. Lower Triassic sequence stratigraphy of the western part of the Germanic Basin (west of Black Forest): Fluvial system evolution through time and space. Sedimentary Geology 186: 187–211.
Broili, F., and E. Fischer. 1917. Trachelosaurus Fischeri nov. gen. nov. sp., ein neuer Saurier aus dem Buntsandstein von Bernburg. Jahrbuch der Königlich Preussischen Geologischen Landesanstalt zu Berlin 37: 359–414.
Butler, R.J., S.L. Brusatte, M. Reich, S.J. Nesbitt, R.R. Schoch, and J.J. Hornung. 2011. The sail-backed reptile Ctenosauriscus from the latest Early Triassic of Germany and the timing and biogeography of the early archosaur radiation. PLoS ONE 6(10): e25693. https://doi.org/10.1371/journal.pone.0025693.
Carroll, R.L. 1976. Noteosuchus—the earliest known rhynchosaur. Annals of the South African Museum 72: 37–57.
Carroll, R.L. 1988. Vertebrate paleontology and evolution. New York: W. H. Freeman & Company.
Chatterjee, S. 1974. A rhynchosaur from the Upper Triassic Maleri formation of India. Philosophical Transactions of the Royal Society of London (B) 267: 209–261.
Cocude-Michel, M. 1963. Les rhynchocéphales et les sauriens des calcaires lithographiques (Jurassique supérieur) d’Europe occidentale. Nouvelles Archives du Muséum d'Histoire Naturelle de Lyon 7: 3–224.
Currie, P.J. 1981. Hovasaurus boulei, an aquatic eosuchian from the Upper Permian of Madagascar. Palaeontologica africana 24: 99–163.
Dilkes, D.W. 1995. The rhynchosaur Howesia browni from the Lower Triassic of South Africa. Palaeontology 38: 665–685.
Dilkes, D.W. 1998. The Early Triassic rhynchosaur Mesosuchus browni and the interrelationships of basal archosauromorph reptiles. Philosophical Transactions of the Royal Society of London (B) 353: 501–541.
Ewer, R.F. 1965. The anatomy of the thecodont reptile Euparkeria capensis Broom. Philosophical Transactions of the Royal Society of London (B) 248: 379–435.
Ezcurra, M.D. 2016. The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. PeerJ 4: e1778.
Ezcurra, M.D., and R.J. Butler. 2018. The rise of the ruling reptiles and ecosystem recovery from the Permian-Triassic mass extinction. Proceedings of the Royal Society of London (B) 285: 20180361. https://doi.org/10.1098/rspb.2018.0361.
Ezcurra, M.D., F. Montefeltro, and R.J. Butler. 2016. The early evolution of rhynchosaurs. Frontiers in Ecology and Evolution 3: 142.
Ezcurra, M.D., L.E. Fiorelli, A.G. Martinelli, S. Rocher, M.B. von Baczko, M. Ezpeleta, J.R.A. Taborda, E.M. Hechenleitner, M.J. Trotteyn, and J.B. Desojo. 2017. Deep faunistic turnovers preceded the rise of dinosaurs in southwestern Pangaea. Nature Ecology and Evolution 1: 1477–1483.
Ezcurra, M.D., S.J. Nesbitt, M. Bronzati, F.M. Dalla Vecchia, F.L. Agnolin, R.B.J. Benson, F. Brissón Egli, S.F. Cabreira, S.W. Evers, A.R. Gentil, R.B. Irmis, A.G. Martinelli, F.E. Novas, L. Roberto da Silva, N.D. Smith, M.R. Stocker, A.H. Turner, and M.C. Langer. 2020. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588: 445–449.
Ezcurra, M.D., S. Bandyopadhyay, and D.J. Gower. 2021a. A new erythrosuchid archosauriform from the Middle Triassic Yerrapalli Formation of South-Central India. Ameghiniana 58: 132–168.
Ezcurra, M.D., L.E. Fiorelli, M.J. Trotteyn, A.G. Martinelli, and J.B. Desojo. 2021b. The rhynchosaur record, including a new stenaulorhynchine taxon, from the Chañares Formation (upper Ladinian–?lowermost Carnian levels) of La Rioja Province, north-western Argentina. Journal of Systematic Palaeontology 18: 1907–1938.
Fraser, N.C. 1988. The osteology and relationships of Clevosaurus (Reptilia: Sphenodontida). Philosophical Transactions of the Royal Society of London (B) 321: 125–178.
Fraser, N.C., and O. Rieppel. 2006. A new protorosaur (Diapsida) from the Upper Buntsandstein of the Black Forest, Germany. Journal of Vertebrate Paleontology 26: 866–871.
Gall, J.-C. 1971. Faunes et paysages du Grès à Voltzia du Nord des Vosges. Essai paléoécologique sur le Buntsandstein supérieur. Mémoires du Service de la Carte Géologique d’alsace et de Lorraine 34: 1–318.
Gall, J.-C., and L. Grauvogel-Stamm. 1999. Paläoökologie des Oberen Buntsandsteins am Westrand des Germanischen Beckens: Der Voltziensandstein im nordöstlichen Frankreich als deltaische Bildung. In Trias—Eine ganz andere Welt. Mitteleuropa im frühen Erdmittelalter, eds. N. Hauschke and V. Wilde, 283–298. Munich: F. Pfeil.
Goloboff, P.A., and S.A. Catalano. 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32: 221–238.
Goloboff, P.A., J.S. Farris, M. Källersjö, B. Oxelman, M.J. Ramírez, and C.A. Szumik. 2003. Improvements to resampling measures of group support. Cladistics 19: 324–332.
Goloboff, P.A., J.S. Farris, and K.C. Nixon. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24: 774–786.
Gottmann-Quesada, A., and P.M. Sander. 2009. A redescription of the early archosauromorph Protorosaurus speneri Meyer, 1832, and its phylogenetic relationships. Palaeontographica (A) 287: 123–220.
Gow, C.E. 1975. The morphology and relationships of Youngina capensis Broom and Prolacerta broomi Parrington. Palaeontologia africana 18: 89–131.
Günther, A. 1867. Contribution to the anatomy of Hatteria (Rhynchocephalus, Owen). Philosophical Transactions of the Royal Society of London 157: 595–629.
Hancox, P.J., J. Neveling, and B.S. Rubidge. 2020. Biostratigraphy of the Cynognathus Assemblage Zone (Beaufort Group, Karoo Supergroup), South Africa. South African Journal of Geology 123: 217–238.
Haque, Z., J.W. Geissman, R.B. Irmis, P.E. Olsen, C. Lepre, H. Buhedma, R. Mundil, W.G. Parker, C. Ramussen, and G.E. Gehrels. 2021. Magnetostratigraphy of the Triassic Moenkopi Formation from the continuous cores recovered in Colorado Plateau Coring Project Phase 1 (CPCP‐1), Petrified Forest National Park, Arizona, USA: correlation of the Early to Middle Triassic strata and biota in Colorado Plateau and its environs. Journal of Geophysical Research Solid Earth. https://doi.org/10.1029/2021JB021899.
Haubold, H. 1971. Die Tetrapodenfährten des Buntsandsteins in der Deutschen Demokratischen Republik und in Westdeutschland und ihre Äquivalente in der gesamten Trias. Paläontologische Abhandlungen (A) 4: 395–548.
Haubold, H. 2006. Die Saurierfährten Chirotherium barthii Kaup, 1835 – das Typusmaterial aus dem Buntsandstein bei Hildburghausen/Thüringen und das “Chirotherium-Monument.” Veröffentlichungen Naturhistorisches Museum Schleusingen 21: 1–100.
Hoffstetter, R., and J.-P. Gasc. 1969. Vertebrae and ribs of modern reptiles. In Biology of the Reptilia, Vol. 1: Morphology A, eds. C. Gans and T.S. Parsons, 201–310. London: Academic Press.
Hone, D.W.E., and M.J. Benton. 2008. A new genus of rhynchosaur from the Middle Triassic of South-West England. Palaeontology 51: 95–115.
Hounslow, M.W., and G. McIntosh. 2003. Magnetostratigraphy of the Sherwood Sandstone Group (Lower and Middle Triassic), south Devon, UK: Detailed correlation of the marine and non-marine Anisian. Palaeogeography, Palaeoclimatology, Palaeoecology 193: 325–348.
Huene, F. von. 1912. Die Cotylosaurier der Trias. Palaeontographica 59: 69–102.
Huene, F. von. 1929. Über Rhynchosaurier und andere Reptilien aus den Gondwana-Ablagerungen Südamerikas. Geologische und Paläontologische Abhandlungen 17: 1–62.
Huene, F. von. 1938. Stenaulorhynchus, ein Rhynchosauride der ostafrikanischen Obertrias. Nova Acta Leopoldina 6: 83–121.
Huene, F. von. 1946. Die großen Stämme der Tetrapoden in den geologischen Zeiten. Biologisches Zentralblatt 65: 268–275.
Huene, F. von. 1956. Paläontologie und Phylogenie der niederen Tetrapoden. Jena: Gustav Fischer.
Jaekel, O. 1904. Über ein neues Reptil aus dem Buntsandstein der Eifel. Zeitschrift der Deutschen Geologischen Gesellschaft 56: 90–94.
Jaekel, O. 1910. Ueber einen neuen Belodonten aus dem Buntsandstein von Bernburg. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 5: 197–229.
Jalil, N.E. 1997. A new prolacertiform diapsid from the Triassic of North Africa and the interrelationships of the Prolacertiformes. Journal of Vertebrate Paleontology 17: 506–525.
Jaquier, V.P., N.C. Fraser, H. Furrer, and T.M. Scheyer. 2017. Osteology of a new specimen of Macrocnemus aff. M. fuyuanensis (Archosauromorpha, Protorosauria) from the Middle Triassic of Europe: potential implications for species recognition and paleogeography of tanystropheid protorosaurs. Frontiers in Earth Science 5: 91. https://doi.org/10.3389/feart.2017.00091.
Koppka, J. 2021. Eifelosaurus—the lonely lizard. Geoconservation Research. https://doi.org/10.30486/gcr.2021.1911759.1035.
Krebs, B. 1969. Ctenosauriscus koeneni (v. Huene), die Pseudosuchia und die Buntsandstein-Reptilien. Eclogae Geologicae Helvetiae 62: 697–714.
Kuhn, O. 1961. Die Familien der rezenten und fossilen Amphibien und Reptilien. Bamberg: Meisenbach.
Langer, M.C., F.C. Montefeltro, D.E. Hone, R. Whatley, and C.L. Schultz. 2010. On Fodonyx spenceri and a new rhynchosaur from the Middle Triassic of Devon. Journal of Vertebrate Paleontology 30: 1884–1888.
Montefeltro, F.C., J.S. Bittencourt, M.C. Langer, and C.L. Schultz. 2013. Postcranial anatomy of the hyperodapedontine rhynchosaur Teyumbaita sulcognathus (Azevedo and Schultz, 1987) from the Late Triassic of southern Brazil. Journal of Vertebrate Paleontology 33: 67–84.
Nosotti, S. 2007. Tanystropheus longobardicus (Reptilia, Protorosauria): Re-interpretations of the anatomy based on new specimens from the Middle Triassic of Besano (Lombardy, northern Italy). Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 35(3): 1–88.
Osborn, H.F. 1903. On the primary division of the Reptilia into two sub-classes, Synapsida and Diapsida. Science 17: 275–276.
Paul, J. 1999. Fazies und Sedimentstrukturen des Buntsandsteins. In Trias—Eine ganz andere Welt Mitteleuropa im frühen Erdmittelalter, eds. N. Hauschke and V. Wilde, 105–114. Munich: F. Pfeil.
Peyer, B. 1937. Die Triasfauna der Tessiner Kalkalpen. XII. Macrocnemus bassanii Nopcsa. Abhandlungen der Schweizerischen Paläontologischen Gesellschaft 59: 1–140.
Pol, D., and I.H. Escapa. 2009. Unstable taxa in cladistic analysis: Identification and the assessment of relevant characters. Cladistics 25: 515–527.
Pritchard, A.C., A.H. Turner, S.J. Nesbitt, R.B. Irmis, and N.D. Smith. 2015. Late Triassic tanystropheids (Reptilia, Archosauromorpha) from northern New Mexico (Petrified Forest Member, Chinle Formation) and the biogeography, functional morphology, and evolution of Tanystropheidae. Journal of Vertebrate Paleontology 35: e911186. https://doi.org/10.1080/02724634.2014.911186.
Rasmussen, C., R. Mundil, R.B. Irmis, D. Geisler, G.E. Gehrels, P.E. Olsen, D.V. Kent, C. Lepre, S.T. Kinney, J.W. Geismann, and W.G. Parker. 2020. U-Pb zircon geochronology and depositional age models for the Upper Triassic Chinle Formation (Petrified Forest National Park, Arizona, USA): Implications for Late Triassic paleoecological and paleoenvironmental change. Geological Society of America Bulletin 133: 539–558.
Rieppel, O. 1989. The hind limb of Macrocnemus bassanii (Nopcsa) (Reptilia: Diapsida): Development and functional anatomy. Journal of Vertebrate Paleontology 9: 373–387.
Röhling, H.-G., and J. Lepper. 2013. Paläogeographie des Mitteleuropäischen Beckens während der tieferen Trias (Buntsandstein). In Stratigraphie von Deutschland XI. Buntsandstein, ed. Deutsche Stratigraphische Kommission, 43–67. Stuttgart: Schweizerbart.
Säilä, L. 2008. The osteology and affinities of Anomoiodon liliensterni, a procolophonid reptile from the Lower Triassic Bundsandstein[sic] of Germany. Journal of Vertebrate Paleontology 28: 1199–1205.
Sander, M., and C.T. Gee. 1994. Der Buntsandstein der Eifel. In Erdgeschichte im Rheinland—Fossilien und Gesteine aus 400 Millionen Jahren, eds. W. von Koenigswald and W. Meyer, 117–124. Munich: F. Pfeil.
Schoch, R.R. 2011. How diverse is the temnospondyl fauna in the Early Triassic of southwestern Germany? Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 261: 49–60.
Schoch, R.R. 2018. The temnospondyl Parotosuchus nasutus (v. Meyer, 1858) from the Early Triassic Middle Buntsandstein of Germany. Palaeodiversity 11: 673–705.
Schoch, R.R. 2019. Osteology of the temnospondyl Trematosaurus brauni Burmeister, 1849 from the Middle Buntsandstein of Bernburg, Germany. Palaeodiversity 12: 41–63.
Spiekman, S.N.F., and T.M. Scheyer. 2019. A taxonomic revision of the genus Tanystropheus (Archosauromorpha Tanystropheidae). Palaeontologia Electronica. https://doi.org/10.26879/1038.
Spielmann, J.A., S.G. Lucas, L.F. Rinehart, and A.B. Heckert. 2008. The Late Triassic archosauromorph Trilophosaurus. New Mexico Museum of Natural History & Science Bulletin 43: 1–177.
Stets, J. 2013. Buntsandstein im Trier-Bitburg-Becken und dessen Umfeld (Südwest-Eifel und West-Hunsrück). In Stratigraphie von Deutschland XI. Buntsandstein, ed. Deutsche Stratigraphische Kommission, 467–486. Stuttgart: Schweizerbart.
Sues, H.-D., and R.R. Reisz. 2008. Anatomy and phylogenetic relationships of Sclerosaurus armatus (Amniota: Parareptilia) from the Buntsandstein (Triassic) of Europe. Journal of Vertebrate Paleontology 28: 1031–1042.
Whatley, R.L. 2005. Phylogenetic relationships of Isalorhynchus genovefae, the rhynchosaur (Reptilia, Archosauromorpha) from Madagascar. Unpublished PhD dissertation, University of California, Santa Barbara.
Zittel, K.A. von. 1911. Grundzüge der Paläontologie (Paläozoologie) (Neubearbeitet von F. Broili, E. Koken und M. Schlosser) II Abteilung Vertebrata. Munich: R. Oldenbourg.
Acknowledgements
We thank Georg Heumann and Martin Sander for the loan of the holotype of Eifelosaurus triadicus. Michael Benton provided helpful comments on a draft of the manuscript. We acknowledge Felipe Montefeltro for allowing us to reproduce a photograph taken by him of the femur of Stenaulorhynchus stockelyi. Fritz Pfeil granted permission to use two illustrations from books published by his company. This research was partially funded by Agencia Nacional de Promoción Científica y Tecnológica (PICT 2018-01186 to MDE). The TNT software is made freely available by the Willi Hennig Society.
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Conflict of interest
The authors declare that they have no competing interests.
Additional information
Handling editor: Jörg Fröbisch.
Appendix
Appendix
Deactivated terminals
Dinocephalosaurus_orientalis, Macrocnemus_obristi, Fuyuansaurus_acutirostris, Pectodens_zhenyuensis, Protanystropheus_antiquus, Trachelosaurus_fischeri, Tanystropheus_haasi, CRILAR_Pv_461, CRILAR_Pv_462, CRILAR_Pv_497, Chanares_rhynchosaur, Eorasaurus_olsoni, Prolacertoides_jimusarensis, Archosaurus_rossicus_holotype, Archosaurus_complete, Panchet_proterosuchid, Vonhuenia_fredericki, C_rossicus_combined, C_magnus_combined, Chasmatosuchus_vjushkovi, Koilamasuchus_gonzalezdiazi, Kalisuchus_rewanensis_holotype, NMQR_3570, Shansisuchus_kuyeheensis, Uralosaurus_combined, Osmolskina_czatkoviensis, Osmolskina_complete, Otter_Sandstone_archosaur, Stagonosuchus_nyassicus, Dagasuchus_santacruzensis, Hypselorhachis_mirabilis, Waldhaus_poposauroid, Vytshegdosuchus_zbeshartensis, Bystrowisuchus_flerovi, Bromsgroveia_walkeri, Moenkopi_poposauroid, Mandasuchus_tanyauchen, Lutungutali_sitwensis, Nyasasaurus_parringtoni.
Characters modified from the character-taxon matrix of Ezcurra et al. (2021a)
472. Pubis, total length (along anterior margin or pubic shaft or tubercle) versus total length of the femur: equal or lower than 0.29 (0); 0.32–0.45 (1); 0.48–0.57 (2); equal or greater than 0.60 (3) (following Ezcurra et al. 2020). We have modified this character because this formulation has proven to be more informative than the original one.
660. Dorsal vertebrae, position of diapophysis in middle and posterior dorsal vertebrae: level with the anterior portion of the centrum (0); level with the anteroposterior middle of the centrum (1). The serial position and the wording of the character was slightly modified here.
Characters added to the data matrix of Ezcurra et al. (2021a)
712. Ilium, pubic peduncle distal articulation: not expanded anteroposteriorly (0); expanded anteroposteriorly (1) (Ezcurra et al. 2020: character 793).
713. Pubis, pubic shaft in anterior or posterior view: straight (0); laterally curved (1) (Ezcurra et al. 2020: character 797).
714. Femur, shaft in lateral or medial view: straight or incipiently posteriorly curved (0); strongly posteriorly curved (New).
715. Femur, apex of the internal trochanter when it is convergent with the proximal end of the bone: at level with or immediately distal to the proximal articular surface of the bone (0); distinctly distal to the proximal articular surface of the bone (1) (New). Inapplicable if the internal trochanter is absent or not convergent with the proximal end of the bone, neither if the fourth trochanter is present.
716. Femur, development of attachment of the caudifemoralis muscle on the ventral/posterior surface of the bone: not distinct or developed as a rugose scar (0); distinct, crest-like, but dorsoventrally/anteroposteriorly lower than the shaft at mid-length (1); distinct, crest-like, dorsoventrally/anteroposteriorly taller than or subequal to the shaft at mid-length (2) ORDERED (modified from Ezcurra et al. 2020: character 803).
Scorings changed in the character-taxon matrix of Ezcurra et al. (2021a).
All taxa.
The scorings of the modified character 472 and characters 712 and 713 follow those in Ezcurra et al. (2020).
Aenigmastropheus parringtoni
Character 660: changed from (0) to (?).
Bentonyx sidensis
Character 279: changed from (2) to (?).
Elorhynchus carrolli
Character 39: changed from (?) to (3).
Eohyosaurus wolvaardti
Character 279: changed from (2) to (?).
Fodonyx spenceri
Character 279: changed form (2) to (1).
Hyperodapedon gordoni
Character 45: changed from (1) to (?).
Character 279: changed from (1) to (0).
Character 482: changed from (0/1) to (1) (ratio = 1.88, Benton 1983: fig. 32d).
Hyperodapedon huxleyi
Character 45: changed from (1) to (?).
Character 279: changed from (1) to (0).
Character 648: changed form (1) to (2).
Isalorhynchus genovefae
Characters 34, 35, 322, 323: changed from (?) to (-).
Character 279: changed from (2) to (0).
Characters 320, 321: changed from (?) to (0).
Jesairosaurus lehmani
Character 324: changed from (0/1) to (1).
Langeronyx brodiei
Character 279: changed from (2) to (?).
Character 614: changed from (?) to (0).
Character 618: changed from (?) to (1).
Noteosuchus colletti
Character 368: changed from (1) to (?).
Rhynchosaurus articeps
Character 279: changed from (2) to (1).
Character 660: changed from (1) to (0) based on SHYMS 4.
Character 675: changed from (?) to (0) based on SHYMS 5.
Stenaulorhynchus stockleyi
Character 322: changed form (1) to (-).
Teyumbaita sulcognathus
Character 45: changed from (1) to (0).
Character 279: changed from (1) to (2).
Character 504: changed from (1) to (0&1).
Trilophosaurus buettneri
Character 458: changed from (2/3) to (1/2).
Ratios:
Eifelosaurus triadicus
Character 377: ca. 1.16–1.40.
Character 472: ca. 0.38.
Rights and permissions
About this article
Cite this article
Sues, HD., Ezcurra, M.D. & Schoch, R.R. Eifelosaurus triadicus Jaekel, 1904, a “forgotten” reptile from the Upper Buntsandstein (Triassic: Anisian) of the Eifel region, Germany. PalZ 96, 275–287 (2022). https://doi.org/10.1007/s12542-021-00584-5
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12542-021-00584-5