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Description of the metoposaurid Anaschisma browni from the New Oxford Formation of Pennsylvania

Published online by Cambridge University Press:  04 May 2021

Bryan M. Gee
Department of Biology, University of Washington, Seattle, WA, USA Department of Ecology & Evolutionary Biology, University of Toronto, Toronto, ON, Canada
Steven E. Jasinski
Section of Paleontology and Geology, State Museum of Pennsylvania, Harrisburg, PA, USA


Metoposaurids are a widespread and ubiquitous constituent of Late Triassic non-marine paleoenvironments. In North America, this group is practically the only large-bodied temnospondyl clade, and is particularly well documented from the American southwest and south-central regions (Arizona, New Mexico, Texas). However, metoposaurids are poorly documented from eastern North America, with fragmentary, doubtfully diagnostic historical material such as “Dictyocephalus elegans” Leidy, 1856 and “Eupelor durus” Cope, 1866. The Zions View (early Norian?) locality in Pennsylvania preserves more-complete material, which previous workers noted as belonging to “Buettneria perfecta” Case, 1922 (=Anaschisma browni Branson, 1905). However, the material has never been described in a fashion that characterizes the anatomy or that justifies the taxonomic assignment, yet it would represent the most complete material in eastern North America and a substantial expansion of this taxon's geographic range. Here we redescribe the Zions View metoposaurid material in detail, differentiating it from Calamops paludosus Sinclair, 1917, the only other Late Triassic temnospondyl from the eastern seaboard, and demonstrating confident affinities with A. browni. Our study is the first to properly justify the taxonomic referral, underscoring the broader importance of proper documentation of voucher specimens, especially for potential geographic outliers. Anaschisma browni is thus the most widely dispersed metoposaurid. Its easternmost documentation underscores the importance of the undersampled and understudied metoposaurid record on the eastern seaboard for understanding the development of a metoposaurid zone of exclusivity in North America and demonstrates the need for further exploration to refine conceptualizations of Late Triassic tetrapod evolution.

Copyright © The Author(s), 2021. Published by Cambridge University Press on behalf of The Paleontological Society

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Antczak, M., and Bodzioch, A, 2018, Ornamentation of dermal bones of Metoposaurus krasiejowensis and its ecological implications: PeerJ, v. 6, p. e5267. ScholarPubMed
Atchley, S.C., Nordt, L.C., Dworkin, S.I., Ramezani, J., Parker, W.G., Ash, S.R., and Bowring, S.A., 2013, A linkage among Pangean tectonism, cyclic alluviation, climate change, and biologic turnover in the Late Triassic: the record from the Chinle Formation, southwestern United States: Journal of Sedimentary Research, v. 83, p. 11471161. Scholar
Baird, D., 1986, Some Upper Triassic reptiles, footprints, and amphibian from New Jersey: The Mosasaur, v. 3, p. 125153.Google Scholar
Baranyi, V., Reichgelt, T., Olsen, P.E., Parker, W.G., and Kürschner, W.M., 2018, N orian vegetation history and related environmental changes: new data from the Chinle Formation, Petrified Forest National Park (Arizona, SW USA): GSA Bulletin, v. 130, p. 775795. Scholar
Barrett, P.M., Sciscio, L., Viglietti, P.A., Broderick, T.J., Suarez, C.A., Sharman, G.R., Jones, A.S., et al. , 2020, The age of the Tashinga Formation (Karoo Supergroup) in the Mid-Zambezi Basin, Zimbabwe and the first phytosaur from mainland sub-Saharan Africa: Gondwana Research, v. 81, p. 445460. Scholar
Bell, C.J., Gauthier, J.A., and Bever, G.S., 2010, Covert biases, circularity, and apomorphies: a critical look at the North American Quaternary Herpetofaunal Stability Hypothesis: Quaternary International, v. 217, p. 3036. Scholar
Benton, M.J., Bernardi, M., and Kinsella, C., 2018, The Carnian Pluvial Episode and the origin of dinosaurs: Journal of the Geological Society, v. 175, p. 10191026. Scholar
Bernardi, M., Gianolla, P., Petti, F.M., Mietto, P., and Benton, M.J., 2018, Dinosaur diversification linked with the Carnian Pluvial Episode: Nature Communications, v. 9, 1499. ScholarPubMed
Bolt, J.R., and Chatterjee, S., 2000, A new temnospondyl amphibian from the Late Triassic of Texas: Journal of Paleontology, v. 74, p. 670683. Scholar
Bonaparte, J.F., 1966, Chronological survey of the tetrapod-bearing Triassic of Argentina: Brevoria, v. 251, p. 113.Google Scholar
Branson, E.B., 1905, Structure and relationships of American Labyrinthodontidae: The Journal of Geology, v. 13, p. 568610.CrossRefGoogle Scholar
Branson, E.B., and Mehl, M. G., 1929, Triassic amphibians from the Rocky Mountain region: The University of Missouri Studies, v. 4, p. 154253.Google Scholar
Brusatte, S.L., Butler, R.J., Mateus, O., and Steyer, J.S., 2015, A new species of Metoposaurus from the Late Triassic of Portugal and comments on the systematics and biogeography of metoposaurid temnospondyls: Journal of Vertebrate Paleontology, v. 35, e912988. Scholar
Brusatte, S.L., Butler, R.J., Niedźwiedzki, G., Sulej, T., Bronowicz, R., and Satkūnas, J., 2013, First record of Mesozoic terrestrial vertebrates from Lithuania: phytosaurs (Diapsida: Archosauriformes) of probable Late Triassic age, with a review of phytosaur biogeography: Geological Magazine, v. 150, p. 110122. Scholar
Buffa, V., Jalil, N.E., and Steyer, J.S., 2019, Redescription of Arganasaurus (Metoposaurus) azerouali (Dutuit) comb. nov. from the Upper Triassic of the Argana Basin (Morocco), and the first phylogenetic analysis of the Metoposauridae (Amphibia, Temnospondyli): Papers in Palaeontology, v. 5, p. 699717. Scholar
Burmeister, K.C., Flynn, J.J., Parrish, J.M., and Wyss, A.R., 2006, Paleogeographic and biostratigraphic implications of new Early Mesozoic vertebrates from Poamay, central Morondava Basin, Madagascar: New Mexico Museum of Natural History and Science Bulletin, v. 37, p. 457475.Google Scholar
Case, E.C., 1922, New reptiles and stegocephalians from the Upper Triassic of western Texas: Carnegie Institute of Washington, v. 321, p. 784.Google Scholar
Case, E.C., 1931, Description of a new species of Buettneria, with a discussion of the brain case: Contributions from the Museum of Paleontology, University of Michigan, v. 3, p. 187206.Google Scholar
Case, E.C., 1932, A collection of stegocephalians from Scurry County, Texas: Contributions from the Museum of Paleontology, University of Michigan, v. 4, p. 156.Google Scholar
Chakravorti, S., and Sengupta, D.P., 2019, Taxonomy, morphometry and morphospace of cranial bones of Panthasaurus gen. nov. maleriensis from the Late Triassic of India: Journal of Iberian Geology, v. 45, p. 317340. Scholar
Chowdhury, T.R., 1965, A new metoposaurid amphibian from the Upper Triassic Maleri Formation of Central India: Philosophical Transactions of the Royal Society of London B, v. 250, p. 152. Scholar
Colbert, E., and Imbrie, J., 1956, Triassic metoposaurid amphibians: Bulletin of the American Museum of Natural History, v. 110, p. 399452.Google Scholar
Cope, E.D., 1866, Observations on extinct vertebrates of the Mesozoic red sandstone: Proceedings of the Academy of Natural Science Philadelphia, v. 18, p. 249250.Google Scholar
Cope, E.D., 1868, Synopsis of the extinct Batrachia of North America: Transactions of the American Philosophical Society, v. 14, p. 208221.Google Scholar
Cornet, B., 1977, The palynostratigraphy and age of the Newark Supergroup [Ph.D. dissertation]: University Park, PA, Pennsylvania State University, 506 p.Google Scholar
Corroy, G., 1928, Les vertébrés du Trias de Lorraine et le Trias Lorrain: Annales de Paleontologie, v. 17, p. 83136.Google Scholar
Cuny, G., and Ramboer, G., 1991, Nouvelles données sur la fauna et l'age de Saint Nicholas de Port: Revue de Paléobiologie, v. 10, p. 6978.Google Scholar
Dal Corso, J., Bernardi, M., Sun, Y., Song, H., Seyfullah, L.J., Preto, N., Gianolla, P., et al. , 2020, Extinction and dawn of the modern world in the Carnian (Late Triassic): Science Advances, v. 6, eaba0099. Scholar
Doyle, K. D., and Sues, H.-D., 1995, Phytosaurs (Reptilia: Archosauria) from the Upper Triassic New Oxford Formation of York County, Pennsylvania: Journal of Vertebrate Paleontology, v. 15, p. 545553. Scholar
Dubiel, R.F., Parrish, J.T., Parrish, J.M., and Good, S.C., 1991, The Pangaean megamonsoon: evidence from the Upper Triassic Chinle Formation, Colorado Plateau: Palaios, v. 6, p. 347370.CrossRefGoogle Scholar
Dubiel, R.F., Hasiotis, S.T., Davidson, S.K., Leleu, S., and North, C.P., 2011, Deposystems, paleosols, and climatic variability in a continental system: the Upper Triassic Chinle Formation, Colorado Plateau, USA, in Davidson, S.K., Leleu, S., and North, C.P., eds., From River to Rock Record: The Preservation of Fluvial Sediments and their Subsequent Interpretation: SEPM (Society for Sedimentary Geology): Special Publication, v. 97, p. 393–421.CrossRefGoogle Scholar
Dunne, E.M., Farnsworth, A., Greene, S.E., Lunt, D.J., and Butler, R.J., 2020, Climatic drivers of latitudinal variation in Late Triassic tetrapod diversity: Palaeontology, v. 64, p. 101117. Scholar
Dutuit, J.M., 1976, Introduction à l’étude paléontologique du Trias continental marocain. Description des premiers stegocephales recueillis dans le couloir d'Argana (Atlas occidental): Memoires du Museum National d'Histoire naturelle, Paris, Series C, v. 36, p. 1253.Google Scholar
Dutuit, J.M., 1978, Description de quelques fragments osseux provenant de la region de Folakara (Trias supérier malgache): Bulletin du Muséum d'Histoire Naturelle, v. 69, p. 7989.Google Scholar
Emmons, E., 1856, Geological Report of the Midland Counties of North Carolina: New York, George P. Putnam and Co., 352 p.Google Scholar
Fortuny, J., Marcé-Nogué, J., de Esteban-Trivigno, S., Gil, L., and Galobart, À., 2011, Temnospondyli bite club: ecomorphological patterns of the most diverse group of early tetrapods: Journal of Evolutionary Biology, v. 24, p. 20402054. ScholarPubMed
Fortuny, J., Marcé-Nogué, J., Gil, L., and Galobart, À., 2012, Skull mechanics and the evolutionary patterns of the otic notch closure in capitosaurs (Amphibia: Temnospondyli): The Anatomical Record, v. 295, p. 11341146. Scholar
Fortuny, J., Marcé-Nogué, J., Steyer, J.S., de Esteban-Trivigno, S., Mujal, E., and Gil, L., 2016, Comparative 3D analyses and palaeoecology of giant early amphibians (Temnospondyli: Stereospondyli): Scientific Reports, v. 6, 30387. Scholar
Fortuny, J., Marcé-Nogué, J., and Konietzko-Meier, D., 2017, Feeding biomechanics of Late Triassic metoposaurids (Amphibia: Temnospondyli): a 3D finite element analysis approach: Journal of Anatomy, v. 230, p. 752765. ScholarPubMed
Fortuny, J., Arbez, T., Mujal, E., and Steyer, J.-S., 2019, Reappraisal of ‘Metoposaurus hoffmani’ Dutuit, 1978, and description of new temnospondyl specimens from the Middle–Late Triassic of Madagascar (Morondava Basin): Journal of Vertebrate Paleontology, v. 39, e1576701. Scholar
Gee, B.M., and Parker, W.G., 2020, Morphological and histological description of small metoposaurids from Petrified Forest National Park, AZ, USA and the taxonomy of Apachesaurus: Historical Biology, v. 32, p. 203233. Scholar
Gee, B.M., Parker, W.G., and Marsh, A.D., 2017, Microanatomy and paleohistology of the intercentra of North American metoposaurids from the Upper Triassic of Petrified Forest National Park (Arizona, USA) with implications for the taxonomy and ontogeny of the group: PeerJ, v. 5, e3183. ScholarPubMed
Gee, B.M., Parker, W.G., and Marsh, A.D., 2020, Redescription of Anaschisma (Temnospondyli: Metoposauridae) from the Late Triassic of Wyoming and the phylogeny of the Metoposauridae: Journal of Systematic Palaeontology, v. 18, p. 233258. Scholar
Gregory, J.T., 1980, The otic notch of metoposaurid labyrinthodonts: in Jacobs, L.L., ed., Aspects of Vertebrate History: Flagstaff, Museum of Northern Arizona Press, p. 125136.Google Scholar
Huber, P., Lucas, S.G., and Hunt, A.P., 1993, Vertebrate biochronology of the Newark Supergroup Triassic, eastern North America: New Mexico Museum of Natural History and Science Bulletin, v. 3, p. 179186.Google Scholar
Hunt, A.P., 1993. Revision of the Metoposauridae (Amphibia: Temnospondyli) and description of a new genus from western North America: Museum of Northern Arizona Bulletin, v. 59, p. 6797.Google Scholar
Hunt, A.P., and Lucas, S.G., 1993, Taxonomy and stratigraphic distribution of late Triassic metoposaurid amphibians from Petrified Forest National Park, Arizona: Journal of the Arizona-Nevada Academy of Science, v. 27, p. 8996.Google Scholar
Jin, C., Dworkin, S., Atchley, S., and Nordt, L., 2018, Eogenetic diagenesis of Chinle sandstones, Petrified Forest National Park (Arizona, USA): a record of Late Triassic climate change: Sedimentology, v. 65, p. 12771300. Scholar
Kent, D.V., Olsen, P.E., and Muttoni, G., 2017, Astrochronostratigraphic polarity time scale (APTS) for the Late Triassic and Early Jurassic from continental sediments and correlation with standard marine stages: Earth-Science Reviews, v. 166, p. 153180. Scholar
Khaldoune, F., Jalil, N.-E., Germain, D., and Steyer, J.-S., 2016, Les vertébrés du Permien et du Trias du Maroc (bassin d'Argana, Haut-Atlas occidental): une fenêtre ouverte sur l'évolution autour de la grande crise fini-paléozoïque: Mémoires de la Société Géologique de France, v. 180, p. 103167.Google Scholar
Kochanov, W.E., and Sullivan, R.M., 1994, Finding phytosaurs in Pennsylvania: the story of Stahle, Sinclair, and Zions View: Pennsylvania Geology, v. 25, p. 38.Google Scholar
Koken, E., 1913, Beiträge zur Kenntnis der Schichten von Heiligkreuz (Abteital, Shdtirol): Abhandlungen der Kaiserlich-Königlichen Geologischen Reichsanstalt, v. 16, p. 143.Google Scholar
Konietzko-Meier, D., and Klein, N., 2013, Unique growth pattern of Metoposaurus diagnosticus krasiejowensis (Amphibia, Temnospondyli) from the Upper Triassic of Krasiejów, Poland: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 370, p. 145157. Scholar
Konietzko-Meier, D., and Sander, P.M., 2013, Long bone histology of Metoposaurus diagnosticus (Temnospondyli) from the Late Triassic of Krasiejów (Poland) and its paleobiological implications: Journal of Vertebrate Paleontology, v. 335, p. 10031018. Scholar
Konietzko-Meier, D., and Wawro, K., 2007, Mandibular dentition in the Late Triassic temnospondyl amphibian Metoposaurus: Acta Palaeontologica Polonica, v. 52, p. 213215.Google Scholar
Konietzko-Meier, D., Gruntmejer, K., Marcé-Nogué, J., Bodzioch, A., and Fortuny, J., 2018, Merging cranial histology and 3D-computational biomechanics: a review of the feeding ecology of a Late Triassic temnospondyl amphibian: PeerJ, v. 6, e4426. ScholarPubMed
Konietzko-Meier, D., Werner, J.D., Wintrich, T., and Sander, P.M., 2019, A large temnospondyl humerus from the Rhaetian (Late Triassic) of Bonenburg (Westphalia, Germany) and its implications for temnospondyl extinction: Journal of Iberian Geology, v. 45, p. 287300. Scholar
Kozur, H.W., and Weems, R.E., 2010, The biostratigraphic importance of conchostracans in the continental Triassic of the northern hemisphere: Geological Society, London, Special Publications, v. 334, p. 315417. Scholar
Kufner, A.M., and Lovelace, D.M., 2018, Taphonomy of two Late Triassic stereospondyl mass death assemblages from the Popo Agie Formation (Fremont County, Wyoming): Society of Vertebrate Paleontology Annual Meeting Program and Abstracts, p. 162.Google Scholar
Lautenschlager, S., Witzmann, F., and Werneburg, I., 2016, Palate anatomy and morphofunctional aspects of interpterygoid vacuities in temnospondyl cranial evolution: The Science of Nature, v. 103, 79. ScholarPubMed
Lehman, J.-P., 1971. Nouveaux vertébrés fossiles du Trias de la série de Zarzaïtine: Annales de Paleontologie, v. 57, p. 71113.Google Scholar
Leidy, J., 1856, Notice of remains of extinct vertebrated animals discovered by Professor E. Emmons: Proceedings of the Academy of Natural Sciences Philadelphia, v. 8, p. 255257.Google Scholar
Leidy, J., 1857, Notice of remains of extinct fishes: Proceedings of the Academy of Natural Sciences Philadelphia, v. 9, p. 167168.Google Scholar
Leonardi, P., 1952, Un osso nasale di Metoposaurus nei tufi a pachicardie dell’ Alpe di Suisi (Dolomiti occidentali): Revista Italiana di Paleontologia, v. 58, p. 145147.Google Scholar
Lindström, S., Irmis, R.B., Whiteside, J.H., Smith, N.D., Nesbitt, S.J., and Turner, A.H., 2016, Palynology of the upper Chinle Formation in northern New Mexico, USA: implications for biostratigraphy and terrestrial ecosystem change during the Late Triassic (Norian–Rhaetian): Review of Palaeobotany and Palynology, v. 225, p. 106131. Scholar
Long, R.A., and Murry, P.A., 1995, Late Triassic (Carnian and Norian) tetrapods from the southwestern United States: New Mexico Museum of Natural History and Science Bulletin, v. 4, p. 1254.Google Scholar
Lucas, S.G., and Sullivan, R.M., 1996, Fossils provide a Pennsylvania standard for part of Late Triassic time: Pennsylvania Geology, v. 27, p. 814.Google Scholar
Lucas, S.G., Heckert, A.B., and Rinehart, L., 2007, A giant skull, ontogenetic variation and taxonomic validity of the Late Triassic phytosaur Parasuchus: New Mexico Museum of Natural History and Science Bulletin, v. 41, p. 222227.Google Scholar
Lucas, S.G., Rinehart, L.F., Heckert, A.B., Hunt, A.P., and Spielmann, J.A., 2016, Rotten Hill: a Late Triassic bonebed in the Texas Panhandle, USA: New Mexico Museum of Natural History and Science Bulletin, v. 72, p. 197.Google Scholar
Lydekker, R. 1885, The Reptilia and Amphibia of the Maleria and Denwa groups: Memoirs of the Geological Survey of India. Palaeontologia Indica, Series IV. Indian Pretertiary Vertebrata, v. 1, Calcutta and London, Geological Survey Office and Trübner & Co., p. 138.Google Scholar
Maisch, M.W., Matzke, A.T., and Sun, G., 2004, A relict trematosauroid (Amphibia: Temnospondyli) from the Middle Jurassic of the Junggar Basin (NW China): Naturwissenschaften, v. 91, p. 589593. Scholar
Mancuso, A.C., Benavente, C.A., Irmis, R.B., and Mundil, R., 2020, Evidence for the Carnian Pluvial Episode in Gondwana: new multiproxy climate records and their bearing on early dinosaur diversification: Gondwana Research, v. 86, p. 104125. Scholar
Martz, J.W., and Parker, W.G., 2017, Revised formulation of the Late Triassic Land Vertebrate “Faunachrons” of western North America: recommendations for codifying nascent systems of vertebrate biochronology, in Ziegler, K.E., and Parker, W.G., eds., Terrestrial Depositional Systems: Amsterdam, Elsevier, p. 39125. Scholar
Milner, A.R., and Schoch, R.R., 2004, The latest metoposaurid amphibians from Europe: Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 232, p. 231252. Scholar
Milner, A.R., Duffin, C., and Delsate, D., 1996, Plagiosaurid and capitosaurid amphibian material from the Late Triassic of Medernach, Grand-Duchy of Luxembourg: preliminary note: Bulletin de la Sociètè belge de Gèologie, v. 104, p. 4253.Google Scholar
Mueller, B.D., 2007, Koskinonodon Branson and Mehl, 1929, a replacement name for the preoccupied temnospondyl Buettneria Case, 1922: Journal of Vertebrate Paleontology, v. 27, p. 225225.CrossRefGoogle Scholar
Mukherjee, D., Ray, S., and Sengupta, D.P., 2010, Preliminary observations on the bone microstructure, growth patterns, and life habits of some Triassic temnospondyls from India: Journal of Vertebrate Paleontology, v. 30, p. 7893. Scholar
Mukherjee, D., Sengupta, D.P., and Rakshit, N., 2020, New biological insights into the Middle Triassic capitosaurs from India as deduced from limb bone anatomy and histology: Papers in Palaeontology, v. 6, p. 93142. Scholar
Muttoni, G., Kent, D.V., Olsen, P.E., Stefano, P.D., Lowrie, W., Bernasconi, S.M., and Hernández, F.M., 2004, Tethyan magnetostratigraphy from Pizzo Mondello (Sicily) and correlation to the Late Triassic Newark astrochronological polarity time scale: Geological Society of America Bulletin, v. 116, p. 10431058. Scholar
Nilsson, T., 1934, Vorläufige mitteilung über einen Stegocephalenfund aus dem Rhät Schonens: Geologiska Föreningens I Stockholm Förhandlingar, v. 56, p. 428442.CrossRefGoogle Scholar
Olsen, P.E., McCune, A.R., and Thomson, K.S., 1982, Correlation of the early Mesozoic Newark Supergroup by vertebrates, principally fishes: American Journal of Science v. 282, p. 1–44. Scholar
Olsen, P.E., Kent, D.V., Cornet, B., Witte, W.K., and Schlische, R.W., 1996, High-resolution stratigraphy of the Newark rift basin (early Mesozoic, eastern North America). Geological Society of America Bulletin, v. 108, p. 4077.<0040:HRSOTN>2.3.CO;2.2.3.CO;2>CrossRefGoogle Scholar
Pardo, J.D., Small, B.J., and Huttenlocker, A.K., 2017, Stem caecilian from the Triassic of Colorado sheds light on the origins of Lissamphibia: Proceedings of the National Academy of Sciences, v. 114, p. E5389–E5395. Scholar
Parker, W.G., and Irmis, R.B., 2005, Advances in Late Triassic vertebrate paleontology based on new material from Petrified Forest National Park, Arizona: New Mexico Museum of Natural History and Science Bulletin, v. 29, p. 4558.Google Scholar
Parker, W.G., and Martz, J.W., 2010, The Late Triassic (Norian) Adamanian–Revueltian tetrapod faunal transition in the Chinle Formation of Petrified Forest National Park, Arizona: Earth and Environmental Science Transactions of the Royal Society of Edinburgh, v. 101, p. 231260. Scholar
Pyron, R.A., and Wiens, J.J., 2013, Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity: Proceedings of the Royal Society B, v. 280, 20131622. ScholarPubMed
Rasmussen, C., Mundil, R., Irmis, R.B., Geisler, D., Gehrels, G.E., Olsen, P.E., Kent, D.V., et al. , 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: GSA Bulletin, v. 133, p. 539558. Scholar
Reichgelt, T., Parker, W.G., Martz, J.W., Conran, J.G., van Konijnenburg-van Cittert, J.H., and Kürschner, W.M., 2013, The palynology of the Sonsela Member (Late Triassic, Norian) at Petrified Forest National Park, Arizona, USA: Review of Palaeobotany and Palynology, v. 189, p. 1828. Scholar
Rinehart, L.F., Lucas, S.G., and Schoch, R.R., 2015, Eocyclotosaurus appetolatus, a new cyclotosaurid amphibian from the Middle Triassic (Perovkan) Moenkopi Formation of New Mexico, USA: Journal of Vertebrate Paleontology, v. 35, e929140. Scholar
Ruffell, A., Simms, M.J., and Wignall, P.B., 2016, The Carnian Humid Episode of the Late Triassic: a review: Geological Magazine, v. 153, p. 271284. Scholar
Sanchez, S., and Schoch, R.R., 2013, Bone histology reveals a high environmental and metabolic plasticity as a successful evolutionary strategy in a long-lived homeostatic Triassic temnospondyl: Evolutionary Biology, v. 40, p. 627647. Scholar
Schaeffer, B., 1967, Late Triassic fishes from the western United States: Bulletin of the American Museum of Natural History, v. 135, p. 285342.Google Scholar
Schoch, R.R., 2000, The status and osteology of two new cyclotosaurid amphibians from the upper Moenkopi Formation of Arizona (Amphibia: Temnospondyli; Middle Triassic): Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen, v. 216, p. 387411. Scholar
Schoch, R.R., 2008, A new stereospondyl from the German Middle Triassic, and the origin of the Metoposauridae: Zoological Journal of the Linnean Society, v. 152, p. 79113. Scholar
Schoch, R.R., and Milner, A.R., 2000, Handbuch Der Paläoherpetologie: Teil 3B: Stereospondyli: München, Verlag Dr. Friedrich Pfeil, 220 p.Google Scholar
Sciscio, L., Viglietti, P.A., Barrett, P.M., Broderick, T.J., Munyikwa, D., Chapelle, K.E., Dollman, K.N., Edwards, S.F., Zondo, M., and Choiniere, J.N., 2020, Sedimentology and palaeontology of the Upper Karoo Group in the Mid-Zambezi Basin, Zimbabwe: new localities and their implications for interbasinal correlation: Geological Magazine, Scholar
Sengupta, D.P., 1988, Chigutisaurid temnospondyls from the Late Triassic of India and review of the family Chigutisauridae: Palaeontology, v. 38, p. 1959.Google Scholar
Sengupta, D.P., 2003, Triassic temnospondyls of the Pranhita-Godavari Basin, India: Journal of Asian Earth Sciences, v. 21, p. 655662. Scholar
Simms, M.J., and Ruffell, A.H., 1989, Synchroneity of climatic change and extinctions in the Late Triassic: Geology, v. 17, p. 265268.<0265:SOCCAE>2.3.CO;2.2.3.CO;2>CrossRefGoogle Scholar
Simms, M.J., and Ruffell, A.H., 1990, Climatic and biotic change in the Late Triassic: Journal of the Geological Society, v. 147, p. 321327. Scholar
Sinclair, W.J., 1917, A new labyrinthodont from the Triassic of Pennsylvania: American Journal of Science, ser. 4, v. 43, p. 319321. Scholar
So, C., Kufner, A.M., Huttenlocker, A.K., Pardo, J., and Lovelace, D., 2018, Unusual fossorial stereospondyl from the Triassic of Wyoming with implications for origins of Gymnophiona: Society of Vertebrate Paleontology Annual Meeting Program and Abstracts, p. 219.Google Scholar
Spielmann, J.A., and Lucas, S.G., 2012, Tetrapod fauna of the Upper Triassic Redonda Formation east-central New Mexico: the characteristic assemblage of the Apachean land-vertebrate faunachron: New Mexico Museum of Natural History and Science Bulletin, v. 55, p. 1119.Google Scholar
Spielmann, J.A., Lucas, S.G., and Heckert, A.B., 2007, Tetrapod fauna of the Upper Triassic (Revueltian) Owl Rock Formation, Chinle Group, Arizona: New Mexico Museum of Natural History and Science Bulletin, v. 41, p. 371383.Google Scholar
Steyer, J.-S., Laurin, M., Castanet, J., and de Ricqlès, A., 2004, First histological and skeletochronological data on temnospondyl growth: palaeoecological and palaeoclimatological implications: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 206, p. 193201. Scholar
Stocker, M.R., Nesbitt, S.J., Kligman, B.T., Paluh, D.J., Marsh, A.D., Blackburn, D.C., and Parker, W.G., 2019, The earliest equatorial record of frogs from the Late Triassic of Arizona: Biology Letters, v. 15, 20180922. ScholarPubMed
Stocker, M.R., Sues, H.-D., and Jasinski, S.E., 2017, Uniting the Upper Triassic deposits of central and north Pangea: the first record of Parasuchus from the Newark Supergroup and its implications for biochronology: Society of Vertebrate Paleontology Annual Meeting Program and Abstracts, p. 199.Google Scholar
Sues, H.-D., and Olsen, P.E., 2015, Stratigraphic and temporal context and faunal diversity of Permian–Jurassic continental tetrapod assemblages from the Fundy rift basin, eastern Canada: Atlantic Geology, v. 51, p. 139205.CrossRefGoogle Scholar
Sues, H.-D., and Schoch, R.R., 2013, Anatomy and phylogenetic relationships of Calamops paludosus (Temnospondyli, Stereospondyli) from the Triassic of the Newark Basin, Pennsylvania: Journal of Vertebrate Paleontology, v. 33, p. 10611070. Scholar
Sulej, T., 2002, Species discrimination of the Late Triassic temnospondyl amphibian Metoposaurus diagnosticus: Acta Palaeontologica Polonica, v. 47, p. 535546.Google Scholar
Sulej, T., 2007, Osteology, variability, and evolution of Metoposaurus, a temnospondyl from the Late Triassic of Poland: Palaeontologica Polonica, v. 64, p. 29139.Google Scholar
Sulej, T., and Majer, D., 2005, The temnospondyl amphibian Cyclotosaurus from the Upper Triassic of Poland: Palaeontology, v. 48, p. 157170. Scholar
Sullivan, R.M., Lucas, S.G., and Randall, K.A., 1995, Late Triassic vertebrate fauna from the Zions View locality (Little Conewago Creek), York County, Pennsylvania: Society of Vertebrate Paleontology Annual Meeting Program and Abstracts, v. 15, p. 55A.Google Scholar
Suteethorn, V., Janvier, P., and Morales, M., 1988, Evidence for a plagiosauroid amphibian in the Upper Triassic Huai Hin Lat Formation: Journal of Southeast Asian Earth Sciences, v. 2, p. 185187. Scholar
Teschner, E.M., Chakravorti, S., Sengupta, D.P., and Konietzko-Meier, D., 2020, Climatic influence on the growth pattern of Panthasaurus maleriensis from the Late Triassic of India deduced from paleohistology: PeerJ, v. 8, e9868. ScholarPubMed
Teschner, E.M., Sander, P.M., and Konietzko-Meier, D., 2018, Variability of growth pattern observed in Metoposaurus krasiejowensis humeri and its biological meaning: Journal of Iberian Geology, v. 44, p. 99111. Scholar
von Huene, F., 1921, Reptilian and stegocephalian remains from the Triassic of Pennsylvania in the Cope collection: Bulletin of the American Museum of Natural History, v. 44, p. 561574.Google Scholar
von Meyer, H., 1842, Labyrinthodonten Genera. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 1842, p. 301304.Google Scholar
von Zittel, K.A., 1888, Handbuch der Paläeontologie. Abteilung 1. Paläozoologie. Band III. Vertebrata (Pisces, Amphibia, Reptilia, Aves): Münich and Leipzig, Oldenbourg, 900 p.Google Scholar
Wanner, H.E., 1926, Some additional faunal remains from the Trias of York County, Pennsylvania: Proceedings of the Academy of Natural Sciences of Philadelphia, v. 78, p. 2128.Google Scholar
Warren, A.A., and Damiani, R., 1999, Stereospondyl amphibians from the Elliott Formation of South Africa: Palaeontologia Africana, v. 35, p. 4554.Google Scholar
Warren, A., and Marsicano, C., 2000, A phylogeny of the Brachyopoidea (Temnospondyli, Stereospondyli): Journal of Vertebrate Paleontology, v. 20, p. 462483.[0462:APOTBT]2.0.CO;2.CrossRefGoogle Scholar
Warren, A., and Snell, N., 1991, The postcranial skeleton of Mesozoic temnospondyl amphibians: a review: Alcheringa, v. 15, p. 4364. Scholar
Warren, A., Rich, T.H., and Vickers-Rich, P., 1997, The last labyrinthodonts: Palaentographica Abteilung A-Stuttgart, v. 247, p. 124.Google Scholar
Watson, D.M.S., 1919, The structure, evolution and origin of the Amphibia—the orders Rhachitomi and Stereospondyli: Philosophical Transactions of the Royal Society of London, Series B, v. 209, p. 173. Scholar
Welles, S.P., and Cosgriff, J., 1965, A revision of the labyrinthodont family Capitosauridae, Moenkopi Formation of northern Arizona: University of California Publications in Geological Sciences, v. 54, p. 1148.Google Scholar
Whiteside, J.H., Lindström, S., Irmis, R.B., Glasspool, I.J., Schaller, M.F., Dunlavey, M., Nesbitt, S.J., Smith, N.D., and Turner, A.H., 2015, Extreme ecosystem instability suppressed tropical dinosaur dominance for 30 million years: Proceedings of the National Academy of Sciences, v. 112, p. 7909–7913. Scholar
Wilson, J.A., 1948. A small amphibian from the Triassic of Howard County, Texas: Journal of Paleontology, v. 22, p. 359361.Google Scholar