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A new mesenosaurine from the lower Permian of Germany and the postcrania of Mesenosaurus: implications for early amniote comparative osteology

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Abstract

Based on an exceptionally well-preserved articulated postcranial skeleton, the early amniote Cabarzia trostheidei gen. et sp. nov. is described. As the lack of the skull hampers its taxonomic assignment, a large sample of basal amniotes is included as part of an exhaustive comparison. Considering the slender, long-limbed proportions of the skeleton, several potential determinations are suggested in order to test for bolosaur, millerettid, araeoscelid, basal neodiapsid, or synapsid affinities. Numerous character conditions are re-evaluated regarding their distributions among early amniote subclades. The closest match to Cabarzia is found to be the middle Permian Mesenosaurus from Russia. The documentation for both genera provides the most complete postcranial descriptions of non-varanodontine varanopids. One of the main differences from Mesenosaurus is the curved ungual phalanx, indicating a use related to predatory behavior. An investigation of limb proportions, such as slender trunks, elongated hindlimbs, and relatively short forelimbs, may suggest the occurrence of facultative bipedalism in Mesenosaurus and Cabarzia. The oldest known mesenosaurine, C. trostheidei from the Asselian/Sakmarian, also pushes back the oldest evidence of bipedal locomotion by more than 15 Ma.

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References

  • Anderson, J.S., and R.R. Reisz. 2004. Pyozia mesenensis, a new, small varanopid (Synapsida, Eupelycosauria) from Russia: “Pelycosaur” diversity in the Middle Permian. Journal of Vertebrate Paleontology 24(1): 173–179.

    Google Scholar 

  • Andreas, D. 1988. The structural dual character of the Rotliegendes in the Thuringian Forest and its surroundings. Zeitschrift für Geologische Wissenschaften 16: 979–992.

    Google Scholar 

  • Andreas, D. 2014. Der Thüringer Wald im Zeitraum der Stefan-Unterperm-Entwicklung-ein Abschnitt der Zentraleuropäischen N-S-Riftzone innerhalb des Mitteleuropäischen Großschollenscharniers. Freiberger Forschungshefte (C: Paläontologie, Stratigraphie, Fazies) 547: 1–181.

    Google Scholar 

  • Andreas, D., and H. Haubold. 1975. Die biostratigraphische Untergliederung des Autun (Unteres Perm) im mittleren Thüringer Wald. Schriftenreihe für geologische Wissenschaften 3: 5–86.

    Google Scholar 

  • Angielczyk, K.D., and L. Schmitz. 2014. Nocturnality in synapsids predates the origin of mammals by over 100 million years. Proceedings of the Royal Society (B: Biological Sciences) 281 (1–8): 20141642. https://doi.org/10.1098/rspb.2014.1642.

    Google Scholar 

  • Barthel, M. 2003. Die Rotliegendflora des Thüringer Waldes, Teil 1: Einführung und Keilblattpflanzen. Veröffentlichungen Naturhistorisches Museum Schleusingen 18: 3–16.

    Google Scholar 

  • Barthel, M., and S. Brauner. 2015. Die Rotliegendflora des Thüringer Waldes. Supplementum. Semana (Veröffentlichungen Naturhistorisches Museum Schleusingen) 30: 3–37.

    Google Scholar 

  • Barthel, M., and R. Rößler. 1995. “Eine gantz unbekanndte Frucht…” – 300 Jahre paläobotanisches Sammeln und Beobachten in Manebach. Veröffentlichungen Naturhistorisches Museum Schleusingen 10: 49–56.

    Google Scholar 

  • Benson, R.B.J. 2012. Interrelationships of basal synapsids: cranial and postcranial morphological partitions suggest different topologies. Journal of Systematic Palaeontology 10: 601–624.

    Google Scholar 

  • Berman, D.S., and R.R. Reisz. 1982. Restudy of Mycterosaurus longiceps (Reptilia, Pelycosauria) from the Lower Permian of Texas. Annals of Carnegie Museum 51: 423–453.

    Google Scholar 

  • Berman, D.S., and S.S. Sumida. 1990. A new species of Limnoscelis (Amphibia, Diadectomorpha) from the Late Pennsylvanian Sangre de Cristo formation of central Colorado. Annals of Carnegie Museum 59(4): 303–341.

    Google Scholar 

  • Berman, D.S., S.S. Sumida, and T. Martens. 1998. Diadectes (Diadectomorpha, Diadectidae) from the Early Permian of central Germany, with description of a new species. Bulletin of the Carnegie Museum of Natural History 67: 53–93.

    Google Scholar 

  • Berman, D.S., A. Henrici, S.S. Sumida, and T. Martens. 2000a. Redescription of Seymouria sanjuanensis (Seymouriamorpha) from the Lower Permian of Germany based on complete, mature specimens with a discussion of paleoecology of the Bromacker locality assemblage. Journal of Vertebrate Paleontology 20: 253–268.

    Google Scholar 

  • Berman, D.S., R.R. Reisz, D. Scott, A. Henrici, S.S. Sumida, and T. Martens. 2000b. Early Permian bipedal reptile. Science 290: 969–972.

    Google Scholar 

  • Berman, D.S., A.C. Henrici, S.S. Sumida, and T. Martens. 2004. New materials of Dimetrodon teutonis (Synapsida: Sphenacodontidae) from the Lower Permian of Germany. Annals of the Carnegie Museum 73(2): 48–56.

    Google Scholar 

  • Berman, D.S., A.C. Henrici, S.S. Sumida, T. Martens, and V. Pelletier. 2014. First European record of a varanodontine (Synapsida: Varanopidae): Member of a unique Early Permian upland paleoecosystem, Tambach Basin, central Germany. In Early Evolutionary History of the Synapsida, eds. C.F. Kammerer, K.D. Angielczyk, and J. Fröbisch, 69–86. New York: Springer. (Vertebrate Paleobiology and Paleoanthropology Series).

    Google Scholar 

  • Bernardi, M., S. Kearns, F. Zorzi, A. Lorenzetti, and M. Fornasiero. 2014. Tridentinosaurus is back. Beginning a complete reexamination of the oldest reptile of the Alps. Giornate di Paleontologia SPI 2014 Abstract volume: 86–87.

  • Bickelmann, C., J. Müller, and R.R. Reisz. 2009. The enigmatic diapsid Acerosodontosaurus piveteaui (Reptilia: Neodiapsida) from the Upper Permian of Madagascar and the paraphyly of ‘‘younginiform’’ reptiles. Canadian Journal of Earth Sciences 46: 651–661.

    Google Scholar 

  • Botha-Brink, J., and S.P. Modesto. 2009. Anatomy and relationships of the Middle Permian Varanopid Heleosaurus scholtzi based on a social aggregation from the Karoo Basin of South Africa. Journal of Vertebrate Paleontology 29(2): 389–400.

    Google Scholar 

  • Boy, J.A., and T. Martens. 1991. Ein neues captorhinomorphes Reptil aus dem thüringischen Rotliegend (Unter-Perm; Ost-Deutschland). Paläontologische Zeitschrift 65: 363–389.

    Google Scholar 

  • Brinkman, D.B., D.S. Berman, and D.A. Eberth. 1984. A new araeoscelid reptile, Zarcasaurus tanyderus, from the Culter Formation (Lower Permian) of north-central New Mexico. New Mexico Geology 6(2): 34–39.

    Google Scholar 

  • Brough, M.C., and J. Brough. 1967. Studies on early tetrapods III. The genus Gephyrostegus. Philosophical Transactions of the Royal Society (B: Biological Sciences) 252: 147–165.

    Google Scholar 

  • Campione, N., and R.R. Reisz. 2010. Varanops brevirostris (Eupelycosauria: Varanopidae) from the Lower Permian of Texas, with discussion of varanopid morphology and interrelationsships. Journal of Vertebrate Paleontology 30(3): 724–746.

  • Carroll, R.L. 1964. The earliest reptiles. Journal of the Linnean Society (Zoology) 45(304): 61–83.

    Google Scholar 

  • Carroll, R.L. 1969. A Middle Pennsylvanian captorhinomorph, and the interrelationships of primitive reptiles. Journal of Paleontology 43(1): 151–170.

    Google Scholar 

  • Carroll, R.R. 1970. The ancestry of reptiles. Philosophical Transactions of the Royal Society (B: Biological Sciences) 257: 267–308.

    Google Scholar 

  • Carroll, R.L. 1976a. Eosuchians and the origin of archosaurs. In Essays on Palaeontology in Honour of Louis Shano Russell, ed. C.S. Churcher, 58–79. Athlon: Royal Ontario Museum. (Life Sciences Miscellaneous Publications).

    Google Scholar 

  • Carroll, R.L. 1976b. Galesphyrus capensis, a younginid eosuchian from the Cistecephalus zone of South Africa. Annals of the South African Museum 72 (4): 59–68.

    Google Scholar 

  • Carroll, R.L. 1977. The origin of lizards. In Problems in Vertebrate Evolution, eds. S.M. Andrews, R.S. Miles, and A.D. Walker, 359–396. London: Academic Press.

    Google Scholar 

  • Carroll, R.L. 1981. Plesiosaur ancestors from the upper permian of Madagascar. Philosophical Transactions of the Royal Society (B: Biological Sciences) 293(1066): 315–383.

    Google Scholar 

  • Carroll, R.L., and D. Baird. 1972. Carboniferous stem-reptiles of the family Romeriidae. Bulletin of the Museum of Comparative Zoology 143(5): 321–363.

    Google Scholar 

  • Carroll, R.L., and P. Gaskill. 1978. The order microsauria. American Philosophical Society Memoirs 126: 1–211.

    Google Scholar 

  • Carroll, R.L., and P. Thompson. 1982. A bipedal lizardlike reptile from the Karroo. Journal of Paleontology 56(1): 1–10.

    Google Scholar 

  • Case, E.C. 1907. Revision of the Pelycosauria of North America. Carnegie Institution of Washington Publications 55: 1–176.

    Google Scholar 

  • Clark, J., and R.L. Carroll. 1973. Romeriid reptiles from the lower Permian. Bulletin of the Museum of Comparative Zoology 144 (5): 353–407.

  • Clemente, C.J. 2014. The evolution of bipedal running in lizards suggests a consequential origin may be exploited in later lineages. Evolution 68(8): 2171–2183.

    Google Scholar 

  • Clemente, C.J., P.C. Withers, G. Thompson, and D. Lloyd. 2008. Why go bipedal? Locomotion and morphology in Australian agamid lizards. Journal of Experimantal Biology 211: 2058–2065.

    Google Scholar 

  • Credner, H. 1889. Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen’schen Grundes bei Dresden. viii. Theil. Kadaliosaurus priscus Crd. Zeitschrift der Deutschen Geologischen Gesellschaft 41: 319–342.

    Google Scholar 

  • Credner, H. 1888. Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen’schen Grundes bei Dresden, vii. Theil: Palaeohatteria longicaudata CRED. Zeitschrift der Deutschen Geologischen Gesellschaft 40: 490–558.

    Google Scholar 

  • Currie, P.J. 1981. Hovasaurus boulei, an aquatic eosuchian from the Upper Permian of Madagascar. Palaeontologia africana 24: 99–168.

    Google Scholar 

  • Currie, P.J. 1982. The osteology and relationships of Tangasaurus mennelli Haughton (Reptilia, Eosuchia). Annals of the South African Museum 86(8): 247–265.

    Google Scholar 

  • Currie, P.J., and R.L. Carroll. 1984. Ontogenetic changes in the eosuchian reptile Thadeosaurus. Journal of Vertebrate Paleontology 4(1): 68–84.

    Google Scholar 

  • Davis, K. 2013. Lower Permian Vertebrates of Oklahoma. Volume 2—Richards Spur. Memphis: D&D Fossils.

  • deBraga, M., and R.R. Reisz. 1995. A new diapsid reptiles from the uppermost Carboniferous (Stephanian) of Kansas. Palaeontology 38(1): 199–212.

    Google Scholar 

  • DeMar, R. 1970. A primitive pelycosaur from the Pennsylvanian of Illinois. Journal of Paleontology 44(1): 154–163.

    Google Scholar 

  • DiMichele, W.A., C.B. Cecil, I.P. Montañez, and H.J. Falcon-Lang. 2010. Cyclic changes in Pennsylvanian paleoclimate and effects on floristic dynamics in tropical Pangaea. International Journal of Coal Geology 83: 329–344.

    Google Scholar 

  • Eberth, D.A., and D. Brinkman. 1983. Ruthiromia elcobriensis, a new pelycosaur from El Cobre Canyon, New Mexico. Breviora 474: 1–26.

    Google Scholar 

  • Eberth, D.A., D.S. Berman, S.S. Sumida, and H. Hopf. 2000. Lower Permian terrestrial paleoenvironment of the Tambach Basin (Thuringia, central Germany): The upland holy grail. Palaios 15: 293–313.

    Google Scholar 

  • Efremov, J.A. 1938. Some new Permian reptiles of the USSR. Proceedings of the USSR Academy of Sciences 19(9): 771–776.

    Google Scholar 

  • Evans, S.E. 1988. The early history and relationships of the Diapsida. In The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds, ed. M.J. Benton, Systematics Association Special Volume 35A: 221–260. Oxford: Clarendon Press.

    Google Scholar 

  • Falconnet, J. 2007. Redescription d’Aphelosaurus lutevensis Gervais, 1859 (Diapsida, Araeoscelidia) du Permien de Lodève. Master thesis. Paris: Université Pierre et Marie Curie, 1–43.

  • Falconnet, J. 2012. First evidence of a bolosaurid parareptile in France (latest Carboniferous–earliest Permian of the Autun basin) and the spatiotemporal distribution of the Bolosauridae. Bulletin de la Société Géologique de France 183(6): 495–508.

    Google Scholar 

  • Falconnet, J. 2013. The Sphenacodontid synapsid Neosaurus cynodus, and related material, from the Permo-Carboniferous of France. Acta Palaeontologica Polonica 60(1): 169–182.

    Google Scholar 

  • Felice, R.N., and K.D. Angielczyk. 2014. Was Ophiacodon (Synapsida, Eupelycosauria) a swimmer? A test using vertebral dimensions. In Early Evolutionary History of the Synapsida, eds. C.F. Kammerer, K.D. Angielczyk, and J. Fröbisch, 25–51. New York: Springer. (Vertebrate Paleobiology and Paleoanthropology Series).

  • Fox, R.C., and M.C. Bowman. 1966. Osteology and relationsships of Captorhinus aguti (COPE) (Reptilia: Captorhinomorpha). The University of Kansas Paleontological Contributions 11: 1–79.

    Google Scholar 

  • Fröbisch, J., R.R. Schoch, J. Müller, T. Schindler, and D. Schweiss. 2011. A new basal sphenacodontid synapsid from the Late Carboniferous of the Saar–Nahe Basin, Germany. Acta Palaeontologica Polonica 56(1): 113–120.

  • Gottmann-Quesada, A., and P.M. Sander. 2009. A redescription of the early archosauromorph Protorosaurus speneri MEYER, 1832, and its phylogenetic relationships. Palaeontographica (A: Paläozoologie und Stratigraphie) 287(4–6): 123–220.

    Google Scholar 

  • Gow, C.E. 1972. The osteology and relationships of the Millerettidae (Reptilia: Cotylosauria). Journal of Zoology 167: 219–264.

    Google Scholar 

  • Gow, C.E. 1975. The morphology and relationships of Youngian capensis Broom and Prolacerta broomi Parrington. Palaeontologia Africana 18: 89–131.

  • Haines, R.W. 1939. A revision of the extensor muscels in the forearm in tetrapods. Journal of Anatomy 73(2): 211–233.

    Google Scholar 

  • Haines, R.W. 1950. The flexor muscles of the forearm and hand in lizards and mammals. Journal of Anatomy 84(1): 13–29.

    Google Scholar 

  • Harris, J.M., and R.L. Carroll. 1977. Kenyasaurus, a new eosuchian reptiles from the early Triassic of Kenya. Journal of Paleontology 51(1): 139–149.

    Google Scholar 

  • Haubold, H. 1970. Versuch der Revision der Amphibien-Fährten des Karbon und Perm. Freiberger Forschungshefte (C: Paläontologie, Stratigraphie, Fazies) 260: 83–117.

    Google Scholar 

  • Haubold, H. 1998. The Early Permian tetrapod ichnofauna of Tambach, the changing concepts in ichnotaxonomy. Hallesches Jahrbuch für Geowissenschaften (B: Geologie, Paläontologie, Mineralogie) 20: 1–16.

    Google Scholar 

  • Haubold, H. 2000. Tetrapodenfährten aus dem Perm—Kenntnisstand und progress 2000. Hallesches Jahrbuch für Geowissenschaften (B: Geologie, Paläontologie, Mineralogie) 22: 1–16.

    Google Scholar 

  • Heaton, M.J., and R.R. Reisz. 1980. A skeletal reconstruction of the Early Permian captorhinid reptile Eocaptorhinus laticeps (Williston). Journal of Paleontology 54(1): 136–143.

    Google Scholar 

  • Heaton, M.J., and R.R. Reisz. 1986. Phylogenetic relationships of captorhinomorph reptiles. Canadian Journal of Earth Sciences 23: 402–418.

    Google Scholar 

  • Henrici, A.C., D.S. Berman, S.G. Lucas, A.B. Heckert, L.F. Rinehart, and K.E. Zeigler. 2005. The carpus and tarsus of the Early Permian synapsid Sphenacodon ferox (Eupelycosauria: Sphenacodontidae). In The Nonmarine Permian, eds. S.G. Lucas, and K.E. Zeigler, New Mexico Museum of Natural History and Science Bulletin 30: 106–110.

    Google Scholar 

  • Heyler, D. 1969. Vertébrés de l’Autunien de France. In Cahiers de Paléontologie, ed. J.P. Lehman, 1–259. Paris: CNRS.

    Google Scholar 

  • Hill, R.V. 2005. Integration of morphological data sets for phylogenetic analysis of Amniota: The importance of integumentary characters and increased taxonomic sampling. Systematic Biology 54 (4): 530–547.

  • Holmes, R.B. 2003. The hind limb of Captorhinus aguti and the step cycle of basal amniotes. Canadian Journal of Earth Sciences 40: 515–526.

    Google Scholar 

  • Hübner, M. 2014. Stratigraphie, Sedimentologie und Paläontologie der Ilmenau- und Goldlauter-Formation im Hartsteinwerk Tabarz, Thüringer Wald. MSc thesis. Freiberg: Technische Universität Bergakademie Freiberg, 1–171.

  • Ivakhnenko, M.F. 2008. Subclass Ophiacomorpha. In Fossil Vertebrates of Russia and Adjacent Countries: Fossil Reptiles and Birds Part 1, eds. M.F. Ivakhnenko and E.N. Kurochkin, 95–100. Moscow: GEOS. (in Russian).

  • Kemp, T.S. 2006. The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis. Journal of Evolutionary Biology 19: 1231–1248.

    Google Scholar 

  • Kim, K.S., J.D. Lim, M.G. Lockley, L. Xing, and Y. Choi. 2017. Korean trackway of a hopping, mammaliform trackmaker is global first from the Cretaceous. Cretaceous Research 74: 188–191. https://doi.org/10.1016/j.cretres.2017.02.005.

    Google Scholar 

  • Kissel, R.A., and R.R. Reisz. 2004. Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes. In Recent Advances in the Origin and Early Radiation of Vertebrates, eds. G. Arratia, M.V.H. Wilson, and R. Cloutier, 409–428. München: Verlag Dr. Friedrich Pfeil.

  • Langston, W. 1965. Oedaleops campi (Reptilia: Pelycosauria), new genus and species from the Lower Permian of New Mexico, and the family Eothyrididae. Bulletin of the Texas Memorial Museum 9: 1–46.

  • Langston, W., and R.R. Reisz. 1981. Aerosaurus wellesi, new species, a varanopseid mammal-like reptile (Synapsida: Pelycosauria) from the Lower Permian of New Mexico. Journal of Vertebrate Paleontology 1(1): 73–96.

  • Laurin, M. 1991. The osteology of a Lower Permian eosuchian from Texas and a review of diapsid phylogeny. Zoological Journal of the Linnean Society 101: 59–95.

    Google Scholar 

  • LeBlanc, A.R.H., and R.R. Reisz. 2014. New postcranial material of the early caseid Casea broilii Williston, 1910 (Synapsida: Caseidae) with a review of the evolution of the sacrum in Paleozoic non-mammalian synapsids. PLoS One 9(12): e115734. https://doi.org/10.1371/journal.pone.0115734.

    Google Scholar 

  • Lee, H.J., Y.N. Lee, A.R. Fiorillo, and J. Lü. 2018. Lizards ran bipedally 110 million years ago. Scientific Reports 8(1): 2617. https://doi.org/10.1038/s41598-018-20809-z.

    Google Scholar 

  • Legler, B., and J.W. Schneider. 2008. Marine ingressions into the Middle/Late Permian saline lake of the Southern Permian Basin (Rotliegend, northern Germany) possibly linked to sea-level highstands in the Arctic rift system. Palaeogeography, Palaeoclimatolology, Palaeoecology 267: 102–114.

    Google Scholar 

  • Leonardi, P. 1959. Tridentinosaurus antiquus Gb. Dal Piaz, rettile protorosauro permiano del Trentino orientale. Memorie degli Instituti di Geologia e Mineralogia dell’Università di Padova XXI: 3–15.

    Google Scholar 

  • Liu, J., and G.S. Bever. 2015. The last diadectomorph sheds light on Late Palaeozoic terapod biogeography. Biology Letters 11: 20150100. https://doi.org/10.1098/rsbl.2015.0100.

    Google Scholar 

  • Lucas, S.G., J. Barrick, K. Krainer, and J.W. Schneider. 2013. The Carboniferous–Permian boundary at Carrizo Arroyo, Central New Mexico, USA. Stratigraphy 10: 153–170.

  • Lützner, H., D. Andreas, J.W. Schneider, S. Voigt, and R. Werneburg. 2012. Stefan und Rotliegend im Thüringer Wald und seiner Umgebung. In Stratigraphie von Deutschland X. Rotliegend. Teil I: Innervariscische Becken, ed. Deutsche Stratigraphische Kommission. Schriftenreihe der Deutschen Gesellschaft für Geowissenschaften 61: 418–487.

    Google Scholar 

  • Maddin, H.C., and R.R. Reisz. 2007. The morphology of the terminal phalanges in Permo-Carboniferous synapsids: An evolutionary perspective. Canadian Journal of Earth Sciences 44: 267–274.

    Google Scholar 

  • Maddin, H.C., D.C. Evans, and R.R. Reisz. 2006. An Early Permian varanodontine varanopid (Synapsida: Eupelycosauria) from the Richards Spur locality, Oklahoma. Journal of Vertebrate Paleontology 26(4): 957–966.

  • Martens, T. 1991. Ein besonderes Fossil. Paläontologische Zeitschrift 65: 225–226.

    Google Scholar 

  • Martens, T. 1992. Untitled. Abhandlungen und Berichte des Museums der Natur Gotha 17: 116.

  • Martens, T. 2010. Ursaurier zwischen Thüringer Wald und Rocky Mountains. Gotha: Museum der Natur Gotha.

    Google Scholar 

  • Martens, T., J.W. Schneider, and H. Walter. 1981. Zur Paläontologie und Genese fossilführender Rotsedimente—Der Tambacher Sandstein, Oberrotliegendes, Thüringer Wald. Freiberger Forschungshefte (C: Paläontologie, Stratigraphie, Fazies) 363: 75–100.

    Google Scholar 

  • Modesto, S.P., and R.R. Reisz. 2002. An enigmatic new diapsid reptile from the Upper Permian of Eastern Europe. Journal of Vertebrate Paleontology 22(4): 851–855.

    Google Scholar 

  • Modesto, S.P., D.M. Scott, and R.R. Reisz. 2009. A new parareptile with temporal fenestration from the Middle Permian of South Africa. Canadian Journal of Earth Sciences 46: 9–20.

    Google Scholar 

  • Modesto, S.P., R.M.H. Smith, N.E. Campione, and R.R. Reisz. 2011. The last “pelycosaur“: A varanopid synapsid from the Pristerognathus Assemblage Zone, Middle Permian of South Africa. Naturwissenschaften 98: 1027–1034.

    Google Scholar 

  • Modesto, S.P., D.M. Scott, M.J. MacDougall, H.-D. Sues, D.C. Evans, and R.R. Reisz. 2015. The oldest parareptile and the early diversification of reptiles. Proceedings of the Royal Society (B: Biological Sciences) 282 (1–9): 20141912. https://doi.org/10.1098/rspb.2014.1912.

    Google Scholar 

  • Montañez, I.P., N.J. Tabor, D. Niemeier, W.A. DiMichele, T.D. Frank, C.R. Fielding, J.L. Isbell, L.P. Birgenheier, and M.C. Rygel. 2007. CO2-forced climate and vegetation instability during Late Paleozoic deglaciation. Science 315: 87–91.

    Google Scholar 

  • Moss, J.L. 1972. The morphology and phylogenetic relationshsips of the Lower Permian tetrapod Tseajaia campi VAUGHN (Amphibia: Seymouriamorpha). University of California Publications in Geological Sciences 98: 1–63.

    Google Scholar 

  • Müller, J., and R.R. Reisz. 2006. The phylogeny of early eureptiles: Comparing parsimony and Bayesian approaches in the investigation of a basal fossil clade. Systematic Biology 55(3): 503–511.

    Google Scholar 

  • Müller, J., D.S. Berman, A.C. Henrici, T. Martens, and S.S. Sumida. 2006. The basal reptile Thuringothyris mahlendorffae (Amniota: Eureptilia) from the Lower Permian of Germany. Journal of Paleontology 80(4): 726–739.

  • Olberding, J.P., L.D. McBrayer, and T.E. Higham. 2012. Performance and three-dimensional kinematics of bipedal lizards during obstacle negotiation. The Journal of Experimental Biology 215: 247–255.

    Google Scholar 

  • Olson, E.C. 1965. New Permian vertebrates from the Chickasha Formation in Oklahoma. Circular Oklahoma Geological Survey 70: 5–70.

  • Olson, E.C. 1970. New and little known genera and species of vertebrates from the Lower Permian of Oklahoma. Fieldiana: Geology 18(3): 359–434.

    Google Scholar 

  • Olson, E.C. 1974. On the source of therapsids. Annals of the South African Museum 64: 27–46.

    Google Scholar 

  • Osborn, H.F. 1903. On the primary division of the Reptilia into two sub-classes, Synapsida and Diapsida. Science 17(424): 275–276.

  • Paton, R.L. 1974. Lower Permian pelycosaurs from the English Midlands. Palaeontology 17(3): 541–552.

    Google Scholar 

  • Peabody, F.E. 1952. Petrolacosaurus kansensis Lane, a Pennsylvanian reptile from Kansas. University of Kansas Paleontological Contributions Article 1: 1–41.

    Google Scholar 

  • Pelletier, V. 2014. Postcranial description and reconstruction of the varanodontine varanopid Aerosaurus wellesi (Synapsida: Eupelycosauria). In Early Evolutionary History of the Synapsida, eds. C.F. Kammerer, K.D. Angielczyk, and J. Fröbisch, 53–68. New York: Springer. (Vertebrate Paleobiology and Paleoanthropology Series).

  • Reisz, R.R. 1972. Pelycosaurian reptiles from the Middle Pennsylvanian of North America. Bulletin of the Museum of Comparative Zoology 144(2): 27–62.

    Google Scholar 

  • Reisz, R.R. 1975. Pennsylvanian pelycosaurs from Linton, Ohio and Nýřany, Czechoslovakia. Journal of Paleontology 49(3): 522–527.

    Google Scholar 

  • Reisz, R.R. 1980. A protorothyridid captorhinomorph reptile from the Lower Permian of Oklahoma. Life Sciences Contributions of the Royal Ontario Museum 121: 1–16.

  • Reisz, R.R. 1981. A diapsid reptile from the Pennsylvanian of Kansas. Special Publications of the Museum of Natural History, University of Kansas 7: 1–74.

  • Reisz, R.R. 1986. Pelycosauria. Handbuch der Paläoherpetologie, Part 17A. Stuttgart: Gustav Fischer Verlag.

    Google Scholar 

  • Reisz, R.R., and D. Baird. 1983. Captorhinomorph “stem” reptiles from the Pennsylvanian coal-swamp deposit of Linton, Ohio. Annals of the Carnegie Museum 52: 393–411.

    Google Scholar 

  • Reisz, R.R., and D.S. Berman. 2001. The skull of Mesenosaurus romeri, a small varanopseid (Synapsida: Eupelycosauria) from the Upper Permian of the Mezen river basin. Northern Russia Annals of Carnegie Museum 70(2): 113–132.

    Google Scholar 

  • Reisz, R.R., and D.W. Dilkes. 2003. Archaeovenator hamiltonensis, a new varanopid (Synapsida: Eupelycosauria) from the Upper Carboniferous of Kansas. Canadian Journal of Earth Sciences 40: 667–678.

    Google Scholar 

  • Reisz, R.R., and J. Fröbisch. 2014. The oldest caseid synapsid from the Late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates. PLoS One 9(4): e94518. https://doi.org/10.1371/journal.pone.0094518.

    Google Scholar 

  • Reisz, R.R., and M. Laurin. 2004. A reevaluation of the enigmatic Permian synapsid Watongia and of its stratigraphic significance. Canadian Journal of Earth Sciences 41: 377–386.

    Google Scholar 

  • Reisz, R.R., and S.P. Modesto. 1996. Archerpeton anthracos from the Joggins Formation of Nova Scotia: a microsaur, not a reptile. Canadian Journal of Earth Sciences 33: 703–709.

    Google Scholar 

  • Reisz, R.R., and S.P. Modesto. 2007. Heleosaurus scholtzi from the Permian of South Africa: a varanopid synapsid, not a diapsid reptile. Journal of Vertebrate Paleontology 27(3): 734–739.

    Google Scholar 

  • Reisz, R.R., and L.A. Tsuji. 2006. An articulated skeleton of Varanops with bite marks: the oldest known evidence of scavenging among terrestrial vertebrate. Journal of Vertebrate Paleontology 26(4): 1021–1023.

    Google Scholar 

  • Reisz, R.R., D.S. Berman, and D. Scott. 1984. The anatomy and relationships of the Lower Permian reptile Araeoscelis. Journal of Vertebrate Paleontology 4(1): 57–67.

    Google Scholar 

  • Reisz, R.R., H. Wilson, and D. Scott. 1997. Varanopseid synapsid skeletal elements from Richards Spur, a Lower Permian fissure fill near Fort Sill, Oklahoma. Oklahoma Geology Notes 57(5): 160–170.

    Google Scholar 

  • Reisz, R.R., D.W. Dilkes, and D.S. Berman. 1998. Anatomy and relationships of Elliotsmithia longiceps Broom, a small synapsid (Eupelycosauria: Varanopseidae) from the Late Permian of South Africa. Journal of Vertebrate Paleontology 18(3): 602–611.

    Google Scholar 

  • Reisz, R.R., S.J. Godfrey, and D. Scott. 2009. Eothyris and Oedaleops: Do these Early Permian synapsids from Texas and New Mexico form a clade? Journal of Vertebrate Paleontology 29(1): 39–47.

    Google Scholar 

  • Reisz, R.R., M. Laurin, and D. Marjanović. 2010. Apsisaurus witteri from the Lower Permian of Texas: yet another small varanopid synapsid, not a diapsid. Journal of Vertebrate Paleontology 30(5): 1628–1631.

    Google Scholar 

  • Reisz, R.R., S.P. Modesto, and D.M. Scott. 2011. A new Early Permian reptile and its significance in early diapsid evolution. Proceedings of the Royal Society (B: Biological Sciences) 278: 3731–3737.

  • Romer, A.S. 1937. New genera and species of pelycosaurian reptiles. Proceedings of the New England Zoological Club 16: 89–96.

    Google Scholar 

  • Romer, A.S. 1976. Osteology of Reptiles. Chicago and London: University of Chicago Press. (3rd impression; original 1956).

  • Romer, A.S., and L.I. Price. 1940. Review of the Pelycosauria. Geological Society of America Special Papers 28: 1–538.

    Google Scholar 

  • Roscher, M., and J.W. Schneider. 2006. Early Pennsylvanian to late Permian climatic development of central Europe in a regional and global context. Geological Society of London, Special Publications 265: 95–136.

    Google Scholar 

  • Ruta, M., J.C. Cisneros, T. Liebrecht, L.A. Tsuji, and J. Müller. 2011. Amniotes through major biological crises: Faunal turnover among parareptiles and the end-Permian mass extinction. Palaeontology 54(5): 1117–1137.

    Google Scholar 

  • Schlotheim, E.F. von. 1804. Beschreibungen merkwürdiger Kräuterabdrücke und Pflanzenversteinerungen. Gotha: Becker.

    Google Scholar 

  • Schneider, J.W. 1994. Environment, biotas and taphonomy of the lacustrine Niederhäslich Limestone, Döhlen Basin, Germany. Transactions of the Royal Society of Edinburgh, Earth Sciences 84: 453–464.

    Google Scholar 

  • Schneider, J.W., and S.G. Lucas. 2015. Late Carboniferous-Permian-Early Triassic nonmarine-marine correlation: Call for global cooperation. Permophiles 61: 28.

  • Schneider, J.W., and R. Romer. 2010. The Late Variscan molasses (Late Carboniferous to Late Permian) of the Saxo-Thuringian Zone. In Pre-Mesozoic Geology of Saxo-Thuringia—From the Cadomian Active Margin to the Variscan Orogen, eds. U. Linnemann, U. Kroner, and R.L. Romer, 323–346. Stuttgart: Schweizerbart, Science Publishers.

  • Schneider, J.W., and F. Scholze. 2016. Late Pennsylvanian–Early Triassic conchostracan biostratigraphy: a preliminary approach. In The Permian Timescale, eds. S.G. Lucas, and S. Shen, Geological Society, London, Special Publications 450. https://doi.org/10.1144/sp450.6.

  • Schneider, J.W., and R. Werneburg. 2012. Biostratigraphie des Rotliegend mit Insekten und Amphibien. In Stratigraphie von Deutschland X. Rotliegend. Teil I: Innervariscische Becken, ed. Deutsche Stratigraphische Kommission. Schriftenreihe der Deutschen Gesellschaft für Geowissenschaften 61: 110–142. Hannover.

  • Schneider, J.W., S.G. Lucas, R. Werneburg, and R. Rößler. 2010. Euramerican Late Pennsylvanian/Early Permian arthropleurid/tetrapod associations—implications for the habitat and paleobiology of the largest terrestrial arthropod. In Carboniferous-Permian Transition in Canon del Cobre, Northern New Mexico, eds. S.G. Lucas, J.W. Schneider, and J.A. Spielmann, New Mexico Museum of Natural History and Science Bulletin 49: 49–70.

  • Schneider, J.W., S.G. Lucas, and J.E. Barrick. 2013. The Early Permian age of the Dunkard Group, Appalachian basin, USA, based on spiloblattinid insect biostratigraphy. International Journal of Coal Geology 119: 88–92.

    Google Scholar 

  • Schneider, J.W., R. Rößler, R. Werneburg, F. Scholze, and S. Voigt. 2014. Part II. The Carboniferous-Permian basins in Saxony, Thuringia, and Saxony-Anhalt of East Germany. In CPC-2014 Field Meeting on Carboniferous and Permian Nonmarine–Marine Correlation. July 21st–27th, Freiberg, Germany. Excursion Guide, eds. J.W. Schneider, S. Oplustil, and F. Scholze, Wissenschaftliche Mitteilungen des Institutes für Geologie, Technische Universität Bergakademie Freiberg 46: 55–121.

  • Schneider, J.W., S. Voigt, S.G. Lucas, and R. Rößler. 2016. Bio- and lithofacies of late Palaeozoic wet red beds and dry red beds. In Fossils: Key to Evolution, Stratigraphy and Palaeoenvironments (87th Annual Conference of the Paläontologische Gesellschaft), eds. B. Niebuhr, M. Wilmsen, L. Kunzmann, and C. Stefen, 1–139. Dresden: Saxoprint.

  • Schneider, J.W., S.G. Lucas, and F. Scholze. 2017. Late Pennsylvanian Blattodea (Insecta) from near Socorro, New Mexico—classification, palecology and biostratigraphy. In Carboniferous-Permian Transition in Socorro County, New Mexico, eds. S.G. Lucas, W.A. DiMichele, and K. Krainer, New Mexico Museum of Natural History and Science Bulletin 77: 5–16.

  • Schoch, R.R., and F. Witzmann. 2009. The temnospondyl Glanochthon from the Lower Permian Meisenheim Formation of Germany. Special Papers in Palaeontology 81: 121–136.

    Google Scholar 

  • Smith, R.M.H., and S.E. Evans. 1996. New material of Youngina: evidence of juvenile aggregation in Permian diapsid reptiles. Palaeontology 39(2): 289–303.

    Google Scholar 

  • Spielmann, J.A., and S.G. Lucas. 2010. Re-evaluation of Ruthiromia elcobriensis (Eupelycosauria: Ophiacodontidae?) from the Lower Permian (Seymourian?) of Cañon del Cobre, Northern New Mexico. In Carboniferous-Permian transition in Cañon del Cobre, eds. S.G. Lucas, J.W. Schneider, and J.A. Spielmann, New Mexico Museum of Natural History and Science Bulletin, 49: 151–158. Albuquerque: New Mexico Museum of Natural History and Science.

    Google Scholar 

  • Spindler, F. 2013. The Niederhäslich tetrapod assemblage (Early Permian, Döhlen basin) from Germany—new insights to ecology, reptiliomorph diversity, and the biology of (basal Sphenacodontia). Society of Vertebrate Paleontology 73rd Annual Meeting Program and Abstracts, p. 218.

  • Spindler, F. 2015. The basal Sphenacodontia—systematic revision and evolutionary implications. PhD thesis. Freiberg: Technische Universität Bergakademie Freiberg, 1–385. http://nbn-resolving.de/urn:nbn:de:bsz:105-qucosa-171748.

  • Spindler, F. 2016. Morphological description and taxonomic status of Palaeohatteria and Pantelosaurus (Synapsida: Sphenacodontia). Freiberger Forschungshefte (C: Paläontologie, Stratigraphie, Fazies) 550(23): 1–57.

    Google Scholar 

  • Spindler, F., J. Falconnet, and J. Fröbisch. 2016. Callibrachion and Datheosaurus, two historical and previously mistaken basal caseasaurian synapsids From Europe. Acta Palaeontologica Polonica 61(3): 597–616.

  • Spindler, F., R. Werneburg, J.W. Schneider, L. Luthardt, V. Annacker, and R. Rößler. 2018. First arboreal ‘pelycosaurs’ (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny. PalZ 92(2): 315–364. https://doi.org/10.1007/s12542-018-0405-9.

  • Stanford, R., M.G. Lockley, C. Tucker, S. Godfrey, and S.M. Stanford. 2018. A diverse mammal-dominated, footprint assemblage from wetland deposits in the Lower Cretaceous of Maryland. Scientific Reports 8(741): 1–12. https://doi.org/10.1038/s41598-017-18619-w.

    Google Scholar 

  • Stein, K., C. Palmer, P.G. Gill, and M.J. Benton. 2008. The aerodynamics of the British Late Triassic Kuehneosauridae. Palaeontology 51(4): 967–981.

  • Steyer, S. 2012. Earth Before the Dinosaurs. Bloomington: Indiana University Press.

  • Sumida, S.S. 1989. Reinterpretation of vertebral structure in the Early Permian pelycosaur Varanosaurus acutirostris (Amniota, Synapsida). Journal of Vertebrate Paleontology 9(4): 451–458.

  • Sumida, S.S. 1997. Locomotor features of taxa spanning the origin of amniotes. In Amniote Origins—Completing the Transition to Land, eds. S.S. Sumida and K.L.M. Martin, 353–398. San Diego: Academic Press.

  • Sumida, S.S., D.S Berman, B. Jefcoat, A. Henrici, and T. Martens. 2013. New Information on the hindlimb structure of the Early Permian bolosaurid reptile Eudibamus cursoris, the earliest known facultative biped. Society of Vertebrate Paleontology 73rd Annual Meeting Program and Abstracts, 222–223.

  • Sumida, S.S., V. Pelletier, and D.S. Berman. 2014. New Information on the basal pelycosaurian-grade synapsid Oedaleops. In Early Evolutionary History of the Synapsida, eds. C.F. Kammerer, K.D. Angielczyk, and J. Fröbisch, 7–23. New York: Springer. (Vertebrate Paleobiology and Paleoanthropology Series).

  • Thommasen, H., and R.L. Carroll. 1981. Broomia, the oldest known millerettid reptile. Palaeontology 24(2): 379–390.

    Google Scholar 

  • Tsuji, L.A. 2010. Evolution, Morphology and Paleobiology of the Pareiasauria and their Relatives (Amniota: Parareptilia). PhD thesis. Berlin: Humboldt-Universität Berlin, 1–220. https://edoc.hu-berlin.de/handle/18452/16918.

  • Tsuji, L.A., J. Müller, and R.R. Reisz. 2012. Anatomy of Emeroleter levis and the phylogeny of the Nycteroleter parareptiles. Journal of Vertebrate Paleontology 32(1): 45–67.

  • Vaughn, P.P. 1955. The Permian reptile Araeoscelis restudied. Bulletin of the Museum of Comparative Zoology 113 (5): 305–467.

    Google Scholar 

  • Voigt, S. 2005. Die Tetrapodenichnofauna des kontinentalen Oberkarbon und Perm im Thüringer Wald—Ichnotaxonomie, Paläoökologie und Biostratigraphie. Göttingen: Cuvillier Verlag.

    Google Scholar 

  • Voigt, S. 2012. Tetrapodenfährten im Rotliegend. In Stratigraphie von Deutschland X. Rotliegend. Teil I: Innervariscische Becken, ed. Deutsche Stratigraphische Kommission. Schriftenreihe der Deutschen Gesellschaft für Geowissenschaften 61: 161–175.

    Google Scholar 

  • Voigt, S., and H. Haubold. 2000. Analyse zur Variabilität der Tetrapodenfährte Ichniotherium cottae aus dem Tambacher Sandstein (Rotliegend, Unterperm, Thüringen). Hallesches Jahrbuch für Geowissenschaften (B: Geologie, Paläontologie, Mineralogie) 22: 17–58.

    Google Scholar 

  • Voigt, S., D.S. Berman, and A.C. Henrici. 2007. First well-established track-trackmaker association of Paleozoic tetrapods based on Ichniotherium trackways and diadectid skeletons from the Lower Permian of Germany. Journal of Vertebrate Paleontology 27: 553–570.

    Google Scholar 

  • Voigt, S., J. Fischer, T. Schindler, M. Wuttke, F. Spindler, and L. Rinehart. 2014. On a potential fossil hotspot for Pennsylvanian-Permian non-aquatic vertebrates in Central Europe. Freiberger Forschungshefte (C: Paläontologie, Stratigraphie, Fazies) 548(22): 39–44.

  • Watson, D.M.S. 1954. On Bolosaurus and the origin and classification of reptiles. Bulletin of the Museum of Comparative Zoology 3 (9): 297–449.

  • Watson, D.M.S. 1957. On Millerosaurus and the early history of the sauropsid reptiles. Philosophical Transactions of the Royal Society (B: Biological Sciences) 240: 325–400.

  • Werneburg, R. 1988. Die Stegocephalen (Amphibia) der Goldlauterer Schichten (Unterrotliegendes, Unterperm) des Thüringer Waldes, Teil III: Apateon dracyiensis (BOY), Branchierpeton reinholdi n. sp. und andere. Veröffentlichungen des Naturkundemuseums Erfurt 7: 80–96.

    Google Scholar 

  • Werneburg, R. 1991. Die Branchiosaurier aus dem Unterrotliegend des Döhlener Beckens bei Dresden. Veröffentlichungen Naturhistorisches Museum Schleusingen 6: 75–99.

    Google Scholar 

  • Werneburg, R. 1999. Ein Pelycosaurier aus dem Rotliegenden des Thüringer Waldes. Veröffentlichungen Naturhistorisches Museum Schleusingen 14: 55–58.

    Google Scholar 

  • Werneburg, R. 2001. Apateon dracyiensis—eine frühe Pionierform der Branchiosaurier aus dem Europäischen Rotliegend, Teil 1: Morphologie. Veröffentlichungen Naturhistorisches Museum Schleusingen 16: 17–36.

    Google Scholar 

  • Werneburg, R. 2002. Apateon dracyiensis—eine frühe Pionierform der Branchiosaurier aus dem Europäischen Rotliegend, Teil 2: Paläoökologie. Veröffentlichungen Naturhistorisches Museum Schleusingen 17: 17–32.

    Google Scholar 

  • Werneburg, R., and S. Brauner. 2014. Der ‘fliegende Branchiosaurier’—eine Eintagsfliege? Freiberger Forschungshefte (C: Paläontologie, Stratigraphie, Fazies) 548(22): 61–69.

  • Werneburg, R., and J.W. Schneider. 2001. Der „Saurierkalkstein“ von Niederhäslich im Döhlen-Becken bei Dresden. In Klassische Fundstellen der Paläontologie, vol. 4, ed. W.K. Weidert, 41–50. Korb: Goldschneck-Verlag.

  • Werneburg, R., and J.W. Schneider. 2006. Amphibian biostratigraphy of the European Permo‐Carboniferous. In Non-Marine Permian Biostratigraphy and Biochronology, eds. S.G. Lucas, G. Cassinis, and J.W. Schneider, Geological Society of London, Special Publications, 265: 201–215.

    Google Scholar 

  • Williston, S.W. 1914. The osteology of some American Permian vertebrates. Journal of Geology 22(4): 364–419.

    Google Scholar 

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Acknowledgments

We owe a great debt of gratitude to Frank Trostheide (Wolmirstedt and Magdeburg) for transporting, loaning, and facilitating the public cataloging of the newly described specimen, as well as for valuable discussions. The documenting carried out in Moscow was greatly facilitated by Ilja Kogan (Freiberg), Evgenia Sytchevskaya, Lev S. Schatenstein, and Valeriy Golubev (Moscow), who provided travel assistance, accommodation, supplies, and collection management. We are grateful for our constructive exchanges with Johannes Müller (Berlin), Michael Buchwitz (Magdeburg), Sebastian Voigt (Thallichtenberg), Thomas Martens (Gotha), and Diane Scott and Robert Reisz (Mississauga). Nico Schendel (Freiberg) kindly assisted with the literature search for the limb plates. Marcel Hübner (Freiberg) and Steffen Trümper (Chemnitz) provided images for the geological map and outcrop documentation, respectively. Stephan Brauner (Friedrichroda) helped by contributing his immense and detailed knowledge regarding local mining history, geology, and re-identifying the horizon of the new holotype. Furthermore, we greatly appreciate the English language help given by Eleonore Horlacher as well as the general funding provided by Michael Völker and Raimund Albersdörfer (Dinosaurier Museum Altmühltal, Denkendorf). The manuscript profited considerably from the kind reviews of Stuart Sumida, Neil Brocklehurst, and one anonymous expert. This research was supported by the Deutsche Forschungsgemeinschaft (DFG grants SCHN 408/12, 20, 21, and 22 to J.W.S.) as well as by the Russian Government as part of the program ‘Competitive Growth of Kazan Federal University Among World’s Leading Academic Centers.’ This work represents a contribution toward the tasks of the IGCP Working Group on Nonmarine-marine late Carboniferous, Permian and early Triassic correlations.

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  1. Dinosaurier-Park Altmühltal, Dinopark 1, 85095, Denkendorf, Germany

    Frederik Spindler

  2. Naturhistorisches Museum Schloss Bertholdsburg, Burgstraße 6, 98553, Schleusingen, Germany

    Ralf Werneburg

  3. TU Bergakademie Freiberg, Geological Institute, Bernhard von Cotta-Straße 2, 09599, Freiberg, Germany

    Jörg W. Schneider

  4. Kazan Federal University, 420008, Kazan, Russia

    Jörg W. Schneider

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  1. Frederik Spindler
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Correspondence to Frederik Spindler.

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Spindler, F., Werneburg, R. & Schneider, J.W. A new mesenosaurine from the lower Permian of Germany and the postcrania of Mesenosaurus: implications for early amniote comparative osteology. PalZ 93, 303–344 (2019). https://doi.org/10.1007/s12542-018-0439-z

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