Saurichthys and other fossil fishes from the late Smithian (Early Triassic) of Bear Lake County (Idaho, USA), with a discussion of saurichthyid palaeogeography and evolution


Authors: Romano C, Kogan I, Jenks J, Jerjen I, Brinkmann W

Published in: Bulletin of Geosciences, volume 87, issue 3; pages: 543 - 570; Received 24 January 2012; Accepted in revised form 21 May 2012; Online 12 September 2012

Keywords: Saurichthyidae, Osteichthyes, coelacanth scale, biotic recovery, biodiversity, cosmopolitanism, extinction, palaeobiogeography, morphological trends,

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A new marine fish assemblage from the late Smithian (Olenekian, Early Triassic) Anasibirites beds of the Thaynes Formation collected near Georgetown (Bear Lake County, south-east Idaho, USA) comprises actinopterygians (Saurichthys cf. elongatus, Actinopterygii indet.), sarcopterygians (Actinistia indet.), and possibly chondrichthyans. We review the global fossil record of the Triassic lower actinopterygian Saurichthys, which is used herein for a case study of trends in morphological adaptations as well as variations in palaeogeographic distribution and diversity dynamics of Early Mesozoic fishes. In the Early Triassic, Saurichthys already occupied a top position in marine food webs, with some species achieving body lengths of up to 1.5 m. Distribution of morphological characters in Saurichthys during the Triassic suggests trends towards a reduction in squamation, stiffening of the fins and axial skeleton, shortening of the postorbital portion of the cranium, and reduction in the number of dermal skull bones. The postcranial adaptations in particular helped to improve the fast-start ability of these ambush predatory fishes. The palaeogeographic range of Saurichthys changed from a virtually global distribution in the Early Triassic (indicating rapid dispersal within marine and freshwater ecosystems after the end-Permian mass extinction) to an occurrence mainly restricted to the north-western Tethys in the Late Triassic, and also towards increasing rarity within continental deposits. Modifications in the palaeogeographic distribution were accompanied by successive loss in global species richness and were possibly related to intra-Triassic extinction events, environmental alterations and/or competition.


Agassiz, L. 1833–1843. Recherches sur les poissons fossiles II, 2. 336 pp. Petitpierre, Neuchatel.

Agassiz, L. 1834. Abgerissene Bemerkungen uber fossile Fische. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und PetREFaktenkunde 1834, 379–390.

Alessandri, G. de 1910. Studii sui pesci Triasici della Lombardia. Memorie della Societa italiana di Scienza naturale 7(1), 1–147.

AREFyev, M.P., Kukhtinov, D.A., Tesakova, E.M. & Shiryaev, A.V. 2009. Development of the ostracod fauna of the Rybinsk Horizon in the Moscow syncline under the conditions of the early Olenekian marine transgression [Razvitiye fauny ostrakod Rybinskogo gorizonta Moskovskoy sineklizy v usloviyah ranneolenekskoy morskoy transgressii], 164–165. In Silantyev, V.V. (ed.) Upper Palaeozoic of Russia: stratigraphy and facies analysis: materials of the Second Russian conference in honor of the 175th birthday of N.A. Golovkinskiy (September 27–30, 2009) [Verkhniy paleozoy Rossii: Stratigrafiya i fatsialnyy analiz: materialy Vtoroy vserossiyskoy konferentsii, posvyashchennoj 175-letiyu so dnya rozhdeniya N.A. Golovkinskogo (27–30 sentyabrya 2009)]. Kazan University, Kazan. [in Russian]

Bachmayer, F. & Warch, A. 1959. Ein bemerkenswerter Fischfund in den Partnachschichten (Trias) sudlich von Feistritz an der Drau (Karnten). Carinthia II 149/69, 42–43.

Bashkuev, A., Sell, J., Aristov, D., Ponomarenko, A., Sinitshenkova, N. & Mahler, H. 2012. Insects from the Buntsandstein of Lower Franconia and Thuringia. Paläontologische Zeitschrift 86(2), 175–185. View article

Bassani, F. 1886. Sui fossili e sull’eta degli schisti bituminosi Triasici di Besano in Lombardia. Communicazione preliminare. Atti della Societa Italiana di Scienze Naturale 29, 15–72.

Bassani, F. 1892. Sui fossili e sull’eta degli schisti bituminosi di Monte Pettine presso Giffoni Valle Piana in Provincia di Salerno (Dolomia Principale). Memoria della Societa Italiana di Scienze, Serie 3 9(3), 1–27.

Bassani, F. 1895. La ittiofauna della Dolomia principale di Giffoni (provincia di Salerno). Paleontographia italica 1, 169–210.

Bellotti, C. 1857. Descrizione di alcune nuove specie di pesci fossili di Perledo e di altre localita lombarde, 419–438. In Stoppani, A. (ed.) Studii Geologici e Paleontologici sulla Lombardia. Turati, Milano.

Bellotti, C. 1873. Osservazioni alla collezione dei pesci fossili del Museo civico. Dal catalogo manoscritto del 1873, 27–30. Reprinted in Pinna, G. 1991. I primi studi sui pesci fossili triassici della Lombardia al Museo di Storia Naturale di Milano. Natura: Rivista di Scienze Naturali 82, 1–41.

Beltan, L. 1968. La faune ichthyologique de l’Eotrias du N.W. de Madagascar: le neurocrâne. 135 pp. Cahiers de paleontologie, CNRS, Paris.

Beltan, L. 1972. La faune ichthyologique du Muschelkalk de la Catalogne. Memorias de la Real Academia de las Ciencias y las Artes de Barcelona 41, 281–325.

Beltan, L. & Janvier, P. 1978. Un nouveau Saurichthyidae (Pisces, Actinopterygii), Saurichthys nepalensis n. sp., du Trias inferieur des Annapurnas (Thakkhola, Nepal) et sa signification paleobiogeographique. Cybium 3(4), 17–28.

Beltan, L., Janvier, P.,Monod, O.& Westphal, F. 1979. A new marine fish and placodont reptile fauna of Ladinian age from southwestern Turkey. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1979(5), 257–267.

Beltan, L. & Tintori, A. 1980. The genus Saurichthys (Pisces, Actinopterygii) during the Gondwana period, 53–59. In Creswell, M.M. & Vella, P. (eds) Gondwana Five, Proceedings of the 5th International Gondwana Symposium, Wellington, New Zealand. A.A. Balkema, Rotterdam.

Bemis, W.E., Findeis, E.K. & Grande, L. 1997. An overview of Acipenseriformes. Environmental Biology of Fishes 48, 25–71. View article

Bender, P.A. 2008. A review of the Early Triassic fish remains from Tasmania. Kanunnah 3, 27–40.

Bender, P.A. & Hancox, P.J. 2004. Newly discovered fish faunas from the Early Triassic, Karoo Basin, South Africa, and their correlative implications. Gondwana Research 7(1), 185–192. View article

Bengtson, P. 1988. Open nomenclature. Palaeontology 31(1), 223–227.

Berg, L.S. 1940. Classification of fishes, both recent and fossil. Travaux de l’Institut de Zoologie de l’Académie des Sciences de l’URSS 5(2), 85–517. [in Russian and English]

Bohm, J. 1903. Uber die obertriadische Fauna der Bareninsel. Kungliga Svenska Vetenskapsakademiens Handlingar 37(3), 2–76.

Brack, P., Rieber, H., Nicora, A. & Mundil, R. 2005. The Global boundary Stratotype Section and Point (GSSP) of the Ladinian stage (Middle Triassic) at Bagolino (Southern Alps, northern Italy) and its implications on the Triassic time scale. Episodes 28(4), 233–244.

Brayard, A., Bucher, H., Escarguel, G., Fluteau, F., Bourquin, S. & Galfetti, T. 2006. The Early Triassic ammonoid recovery: Paleoclimatic significance of diversity gradients. Palaeogeography, Palaeoclimatology, Palaeoecology 239, 347–395. View article

Brinkmann, W., Romano, C., Bucher, H., Ware, D. & Jenks, J. 2010. Palaeobiogeography and stratigraphy of advanced gnathostomian fishes (Chondrichthyes and Osteichthyes) in the Early Triassic and from selected Anisian localities (Report 1863–2009). Zentralblatt für Geologie und Paläontologie, Teil 2 2009(5/6), 765–812.

Bronn, H.G. 1858. Beitrage zur triassischen Fauna und Flora der bituminosen Schiefer von Raibl. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und PetREFaktenkunde 1858(1), 1–32.

Bruhwiler, T., Bucher, H., Brayard, A. & Goudemand, N. 2010. High-resolution biochronology and diversity dynamics of the Early Triassic ammonoid recovery: the Smithian faunas of the Northern Indian Margin. Palaeogeography, Palaeoclimatology, Palaeoecology 297(2), 491–501. View article

Bucher, H. 1989. Lower Anisian ammonoids from the northern Humboldt Range (northwestern Nevada, USA) and their bearing upon the Lower–Middle Triassic boundary. Eclogae Geologicae Helvetiae 82, 945–1002.

Burgin, T. 1990. Reproduction in Middle Triassic actinopterygians; complex fin structures and evidences of viviparity in fossil fishes. Zoological Journal of the Linnean Society 100, 379–391. View article

Burgin, T., Eichenberger, U., Furrer, H. & Tschanz, K. 1991. Die Prosanto Formation – eine fischreiche Fossil-Lagerstatte in der Mitteltrias der Silvretta-Decke (Kanton Graubunden, Schweiz). Eclogae Geologicae Helvetiae 84(3), 921–990.

Cartanya, J. 1999. An overview of the Middle Triassic actinopterygians from Alcover, Mont-ral and El Pinetell (Catalonia, Spain), 535–551. In Arratia, G. & Schultze, H.-P. (eds) Mesozoic Fishes 2. Systematics and Fossil Record. Dr. Friedrich Pfeil, Munchen.

Chang, M.-M. & Miao, D. 2004. An overview of Mesozoic fishes in Asia, 535–563. In Arratia, G. & Tintori, A. (eds) Mesozoic Fishes 3. Systematics, Palaeoenvironments and Biodiversity. Dr. Friedrich Pfeil, Munchen.

Chou, H.-H. & Liu, H.-T. 1957. Fossil fishes from Huanshan, Shensi. Acta Palaeontologica Sinica 5(2), 295–305.

Coates, M.I. 1999. Endocranial preservation of a Carboniferous actinopterygian from Lancashire, UK, and the interrelationships of primitive actinopterygians. Philosophical Transactions of the Royal Society of London B 354, 435–462. View article

Cope, E.D. 1871. Contribution to the ichthyology of the Lesser Antilles. Transactions of the American Philosophical Society 14, 445–483. View article

Cope, E.D. 1872. Observations on the systematic relations of the fishes. Proceedings of the American Association for the Advancement of Science 20, 317–343.

Cope, E.D. 1887. Geology and palaeontology. Zittel’s Manual of Palaeontology. American Naturalist 22(11), 1014–1019.

Costa, O.G. 1862. Studii sopra i terreni ad ittiolitti del Regno di Napoli diretti a stabilire l’eta geologica dei medesimi. Parte I: Scisti bituminiferi di Giffoni. Estratta dall’appendice agli atti della Real Accademia delle Scienze 1862, 1–44.

Deecke, W. 1889. Ueber Fische aus verschiedenen Horizonten der Trias. Palaeontographica 35, 97–138.

Detterman, R.L., Reiser, H.N., Brosge, W.P. & Dutro, J.T. Jr. 1975. Post-Carboniferous stratigraphy, northeastern Alaska. U.S. Geological Survey, Professional Paper 886, 1–46.

Dziewa, T.J. 1980. Early Triassic osteichthyans from the Knocklofty Formation of Tasmania. Papers and Proceedings of the Royal Society of Tasmania 114, 145–160.

Eck, H. 1865. Ueber die Formationen des bunten Sandsteins und des Muschelkalks in Oberschlesien und ihre Versteinerungen. 148 pp. J.F. Stracke, Berlin.

Embry, A.F. 1997. Global sequence boundaries of the Triassic and their identification in the Western Canada Sedimentary Basin. Bulletin of Canadian Petroleum Geology 45(4), 415–433.

Embry, A.F., Krajewski, K.P. & Mork, A. 2002. A Triassic upwelling zone: the Shublik Formation, Arctic Alaska, U.S.A. – Discussion. Journal of Sedimentary Research 72(5), 740–741. View article

Emmert, U. 1977. Geologische Karte von Bayern 1:25000. Erläuterungen zum Blatt Nr. 6035 Bayreuth. 180 pp. Bayerisches Geologisches Landesamt, Munchen.

Evans, H.M. 1904. A new cestraciont spine from the Lower Triassic of Idaho. University of California Publications – Bulletin of the Department of Geology 3, 397–402.

Feldkamp, L.A., Davis, L.C. & Kress, J.W. 1984. Practical cone-beam algorithm. Journal of the Optical Society of America A 1, 612–619. View article

Firtion, F. 1934. Description d’une nouvelle espece de Saurichthys du Gres a Voltzia de Wasselonne. Bulletin du Service de la Carte géologique d’Alsace et de Lorraine 2, 89–97.

Firtion, F. & Schneider, E. 1975. Acidorhynchus im Unteren Lias der Zaberner Senke. Annales universitatis Saraviensis, Mathematisch-Naturwissenschaftliche Fakultät 12, 7–12.

Forey, P.L. 1998. History of the coelacanth fishes. 419 pp. Chapman & Hall, London.

Fortuny, J., Bolet, A., Selles, A.G., Cartanya, J. & Galobart, A. 2011. New insights on the Permian and Triassic vertebrates from the Iberian Peninsula with emphasis on the Pyrenean and Catalonian basins. Journal of Iberian Geology 37(1), 65–86. View article

Frech, F. 1903–1908. Lethaea geognostica 2, Das Mesozoicum, 1. Trias. 623 pp. Schweitzerbart’sche Verlagsbuchhandlung, Stuttgart.

Friedman, M. & Sallan, L.C. 2012. Five hundred million years of extinction and recovery: a Phanerozoic survey of large-scale diversity patterns in fishes. Palaeontology 55(4), 707–742. View article

Furrer, H. 2009. The diet of the predator fish Saurichthys from two Middle Triassic Plattenkalk lagerstatten in the Swiss Alps. 5th International Symposium on Lithographic Limestone and Plattenkalk, Basel, Abstracts and Field Guides, 30–31.

Galfetti, T., Hochuli, P.A., Brayard, A., Bucher, H., Weissert, H. & Vigran, J.O. 2007. Smithian-Spathian boundary event: evidence for global climatic change in the wake of the end-Permian biotic crisis. Geology 35, 291–294. View article

Gardiner, B.G. 1960. A revision of certain actinopterygian and coelacanth fishes, chiefly from the Lower Lias. Bulletin of the British Museum (Natural History), Geology 4(7), 239–384.

Gardiner, B.G. & Schaeffer, B. 1989. Interrelationships of lower actinopterygian fishes. Zoological Journal of the Linnean Society 97, 135–187. View article

Gardiner, B.G., Schaeffer, B. & Masserie, J.A. 2005. A review of lower actinopterygian phylogeny. Zoological Journal of the Linnean Society 144, 511–525. View article

Giraud, M.M., Castanet, J., Meunier, F.J. & Bouligand, Y. 1978. The fibrous structure of coelacanth scales: a twisted ‘plywood’. Tissue and Cell 10(4), 671–686. View article

Griffith, J. 1959. On the anatomy of two saurichthyid fishes, Saurichthys striolatus (Bronn) and S. curionii (Bellotti). Proceedings of the Zoological Society of London 132, 587–606. View article

Griffith, J. 1962. The Triassic fish Saurichthys krambergeri Schlosser. Palaeontology 5(2), 344–354.

Griffith, J. 1977. The Upper Triassic fishes from Polzberg bei Lunz, Austria. Zoological Journal of the Linnean Society 60, 1–93. View article

Griffith, J. 1978. A fragmentary specimen of Saurichthys sp. from the Upper Beaufort series of South Africa. Annals of the South African Museum 76(8), 299–307.

Guex, J., Hungerbuhler, A., Jenks, J.F., O’Dogherty, L., Atudorei, V., Taylor, D.G., Bucher, H. & Bartolini, A. 2010. Spathian (Lower Triassic) ammonoids from western USA (Idaho, California, Utah and Nevada). Mémoirs de Géologie (Lausanne) 49, 1–92.

Hagdorn, H. & Mutter, R.J. 2011. The vertebrate fauna of the Lower Keuper Albertibank (Erfurt Formation, Middle Triassic) in the vicinity of Schwabisch Hall (Baden-Wurttemberg, Germany). Palaeodiversity 4, 223–243.

Hammer, O., Harper, D.A.T. & Ryan, P.D. 2001. Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4(1), 1–9.

Hauff, B. 1938. Uber Acidorhynchus aus den Posidonienschiefern von Holzmaden. Paläontologische Zeitschrift 20, 214–248.View article

Hauff, B. 1938. Uber Acidorhynchus aus den Posidonienschiefern von Holzmaden. Paläontologische Zeitschrift 20, 214–248.

Hennig, E. 1909. Saurichthys-Funde von Rudersdorf. Centralblatt für Mineralogie, Geologie und Paläontologie 1909, 54–60.

Hilzheimer, M. 1930. Ein Saurichthys Ag. aus dem Rot von Rudersdorf. Brandenburgia. Monatsblatt der Gesellschaft für Heimatkunde und Heimatschutz in der Mark Brandenburg 39, 18–20.

Hitij, T., Tintori, A., Žalohar, J., Renesto, S., Celarc, B., Križnar, M. & Kolar-Jurkovšek, T. 2010. New fossil sites with Triassic vertebrate fauna from the Kamnik-Savinja Alps, Slovenia. International Symposium on Triassic and Later Marine Vertebrate Faunas, Beijing. Program and Abstract, 42–46.

Huxley, T.H. 1880. On the application of the laws of evolution to the arrangement of the Vertebrata, and more particularly of the Mammalia. Proceedings of the Scientific Meetings of the Zoological Society of London 1880, 649–662.

Jain, S.L. 1984. A new Triassic fish (Actinopterygii; Saurichthiformes) from the Yerrapalli Formation, Pranhita-Godavari Valley, India. Journal of the Geological Society of India 25(9), 604–610.

Jenkins, F.A. Jr., Shubin, N.H., Amaral, W.W., Gatesy, S.M., Schaff, C.R., Clemmensen, L.B., Downs, W.R., Davidson, A.R., Bonde, N. & Osback, F. 1994. Late Triassic continental vertebrates and depositional environments of the Fleming Fjord Formation, Jameson Land, East Greenland. Meddelelser om Gronland, Geoscience 32, 1–25.

Jenks, J. in press. Ammonoid biostratigraphy of the early Spathian Columbites parisianus Zone (Early Triassic) at Bear Lake Hot Springs, Idaho. The Global Triassic 2. New Mexico Museum of Natural History and Science Bulletin.

Jin, F. 2006. An overview of Triassic fishes from China. Vertebrata PalAsiatica 44(1), 28–42.

Kak, A.C. & Slaney, M. 1988. Algorithms for reconstruction with non-diffracting sources, 49–112. In Cotellessa, R.F. (ed.) Principles of Computerized Tomographic Imaging. IEEE Press, New York.

Kear, B.P., Rich, T.H., Vickers-Rich, P., Ali, M.A., Al-Mufarrih, Y.A.,Matari, A.H., Al-Masary, A.M. & Halawani, M.A. 2010. A review of aquatic vertebrate remains from the Middle–Upper Triassic Jilh Formation of Saudi Arabia. Proceedings of the Royal Society of Victoria 122(1), 1–8.

Khorana, R.K. & Tirkey, B. 1977. Fossil fish Saurichthys from the Upper Triassic of Sonamarg Tilel area, North Kashmir, Jammu and Kashmir. Bulletin of the Indian Geologists Association 10(2), 57–59.

Kner, R. 1866. Die Fische der bituminosen Schiefer von Raibl in Karnthen. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse 53, 152–197.

Kner, R. 1867. Nachtrag zur fossilen Fauna der Asphaltschiefer von Seefeld in Tirol. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse 56(1), 898–909.

Kogan, I. 2011. Remains of Saurichthys (Pisces, Actinopterygii) from the Early Triassic Wordie Creek Formation of East Greenland. Bulletin of the Geological Society of Denmark 59, 93–100.

Kogan, I., Schonberger, K., Fischer, J. & Voigt, S. 2009. A nearly complete skeleton of Saurichthys orientalis (Pisces, Actinopterygii) from the Madygen Formation (Middle to Late Triassic, Kyrgyzstan, Central Asia) – preliminary results. Paläontologie, Stratigraphie, Fazies 17, Freiberger Forschungshefte C 532, 139–152.

Kummel, B. Jr. 1943. The Thaynes Formation, Bear Lake Valley, Idaho. American Journal of Science 241, 316–332. View article

Kummel, B. Jr. 1954. Triassic stratigraphy of southeastern Idaho and adjacent areas. U.S. Geological Survey Professional Paper 254-H, 165–194.

Lambe, L.M. 1914. Description of a new species of Platysomus from the neighborhood of Banff, Alberta. Transactions of the Royal Society of Canada, Series 3, Section 4 8, 17–23.

Lehman, J.-P. 1952. Etude complementaire des poissons de l’Eotrias de Madagascar. Kungliga Svenska Vetenskapsakademiens Handlingar 4(2), 1–192.

Lehman, J.-P. 1964. Etude d’un Saurichthyde de la region d’Oden (Espagne). Annales de Paléontologie (Vertébrés) 50(1), 25–30.

Lehman, J.-P., Chateau, C., Laurain, M. & Nauche, M. 1959. Paleontologie de Madagascar XXVII, Les poissons de la Sakamena moyenne. Annales de Paléontologie 45, 175–219.

Liu, G.B., Zhu, Z.X., Zhang, X.L. & Ai, F. 1999. A coelacanth fossil from Huachi area, Gansu Province. Geological Journal of China Universities 5, 474–480. [in Chinese with English abstract]

Liu, H.-T. 1964. A new coelacanth from the marine Lower Triassic of N.W. Kwangsi, China. Vertebrata PalAsiatica 2(2–3), 147–150.

Liu, X. & Wei, F. 1988. A new saurichthyid from the Upper Permian of Zhejiang, China. Vertebrata PalAsiatica 26(4), 77–89.

Lombardo, C. & Tintori, A. 2005. Feeding specializations in Late Triassic fishes. Annali dell’Universita degli Studi di Ferrara. Museologia Scientifica e Naturalistica, volume speciale 2005, 25–32.

Lombardo, C., Sun, Z.Y., Tintori, A., Jiang, D.Y. & Hao, W.C. 2011. A new species of Perleidus (Actinopterygii) from the Middle Triassic of Southern China. Bollettino della Societa Paleontologica Italiana 50(2), 75–83. View article

Lopez-Arbarello, A. 2004. The record of Mesozoic fishes from Gondwana (excluding India and Madagascar), 597–624. In Arratia, G. & Tintori, A. (eds) Mesozoic fishes 3. Systematics, Paleoenvironments and Biodiversity. Dr. Friedrich Pfeil, Munchen.

Lozovsky, V.R. 1987. Triassic lakes of the Moscow and the Polish-Lithuanian synclines [Triasovy’e ozera Moskovskoy i Polsko-Litovskoy sinekliz.], 199–213. In Martinson, G.G. & Neustruyeva, I.Y. (eds) History of the lakes of Late Palaeozoic and Early Mesozoic [Istoriya ozer posdnego paleozoya i rannego mesozoya]. Nauka, Leningrad. [in Russian]

Maisey, J.G. 1977. The fossil selachian fishes Palaeospinax Egerton, 1872 and Nemacanthus Agassiz, 1837. Zoological Journal of the Linnean Society 60, 259–273. View article

Martin, K. 1873. Ein Beitrag zur Kenntniss fossiler Euganoiden. Zeitschrift der Deutschen Geologischen Gesellschaft 25, 699–735.

Martin, M., Cuny, G. & Mazin, J.-M. 1991. Rostres de Saurichthys (Saurichthyiformes, Saurichthyidae) en trois dimensions dans le Trias moyen d’Israel. Geobios 24(2), 183–186. View article

Massare, J.A. & Callaway, J.M. 1994. Cymbospondylus (Ichthyosauria: Shastasauridae) from the Lower Triassic Thaynes Formation of southeastern Idaho. Journal of Vertebrate Paleontology 14(1), 139–141. View article

Meunier, F.J. 1987. Os cellulaire, os acellulaire et tissus derives chez les Osteichtyens: les phenomenes de l’acellularisation et de la perte de mineralisation. L’Année Biologique 26, 201–233.

Meunier, F.J., Erdmann, M.V., Fermon, Y. & Caldwell, R.L. 2008. Can the comparative study of the morphology and histology of the scales of Latimeria menadoensis and L. chalumnae (Sarcopterygii: Actinistia, Coelacanthidae) bring new insight on the taxonomy and biogeography of recent coelacanthids?, 351–360. In Cavin, L., Longbottom, A. & Richter, M. (eds) Fishes and the break-up of Pangaea. Geological Society, London, Special Publications 295. View article

Meyer, H. von 1849. Fossile Fische aus dem Muschelkalk von Jena, Querfurt und Esperstadt. Palaeontographica 1(5), 195–208.

Miller, W.A. 1979. Observations on the structure of mineralized tissues of the coelacanth, including the scales and their associated odontodes, 68–78. In McCosker, J.E. & Lagios, M.D. (eds) The biology and physiology of the living coelacanth. Occasional Papers of the California Academy of Sciences 134.

Milner, A.R.C., Kirkland, J.I. & Birthisel, T.A. 2006. The geographic distribution and biostratigraphy of Late Triassic-Early Jurassic freshwater fish faunas of the southwestern United States, 522–529. In Harris, J.D., Lucas, S.G., Spielmann, J.A., Lockley, M.G., Milner, A.R.C. & Kirkland, J.I. (eds) The Triassic-Jurassic Terrestrial Transition. New Mexico Museum of Natural History and Science Bulletin 37.

Minikh, A.V. 1981. Saurichthys from the Triassic of the USSR [Zaurikhtisy iz triasa SSSR]. Paleontologicheskiy Zhurnal 1981(1), 105–113. [in Russian]

Minikh, A.V. 1982. New species of Saurichthys from the Early Triassic from the area along the upper Volga [Novye vidy zaurichtisov iz rannego triasa verkhnego Povolzhya]. Ezhegodnik Vsesoyuzhnogo Paleontologicheskogo Obshchestva 25, 205–213. [in Russian]

Minikh, A.V. 1983. Distribution of saurichthyid remains in the Triassic of the eastern European part of the USSR [Raspredeleniye ostatkov zaurikhtiid v triase vostoka Evropeyskoy chasti SSSR], 84–90. In Vostryakov, A.V. (ed.) Geology of the Southern Ural and Volga region. Geophysics, geochemistry, palaeontology [Geologiya Yuzhnogo Urala i Povolzh’ya. Geofizika, geokhimiya, paleontologiya]. Saratov State University Press, Saratov. [in Russian]

Minikh, A.V. 1992. Saurichthys (Order Saurichthyiformes) from the Middle Triassic of the southern Cis-Ural region [Zaurikhtisy (otryad Saurichthyiformes) iz srednego triasa yuzhnogo Priural’ya]. Voprosy Paleontologii 10, 146–151. [in Russian]

Minikh, A.V. & Minikh, M.G. 2005. Fishes of Eastern Europe at the Palaeozoic-Mesozoic boundary. Trophic relationships and palaeogeographical aspects [Ryby Vostochnoy Evropy na rubezhe paleozoya i mesozoya. Troficheskiye svyazi i paleogeograficheskiye aspekty], 35–37. In Podobina, V.M. (ed.) Evolution of the Life on Earth: Materials of the 3rd International Symposium. November 1–3, 2005, Tomsk [Evolutsiya zhizhni na Zemle: Materialy 3-go Mezhdunarodnogo Simposiuma, 1–3 noyabrya 2005 goda, Tomsk]. Tomsk University, Tomsk. [in Russian]

Mork, A., VIGRAN, J.O. & HOCHULI, P.A. 1990. Geology and palynology of the Triassic succession of Bjornoya. Polar Research 8, 141–163.

Mundil, R., Ludwig, K.R., Metcalfe, I. & Renne, P.R. 2004. Age and timing of the Permian mass extinctions: U/Pb dating of closed-system zircons. Science 305, 1760–1763. View article

Munster, G. zu 1839. Ueber einige merkwurdige Fische aus dem Kupferschiefer und dem Muschelkalk. Beiträge zur PetREFacten-Kunde 1, 114–121.

Mutter, R.J., Cartanya, J. & Basaraba, S.A.U. 2008. New evidence of Saurichthys from the Lower Triassic with an evaluation of early saurichthyid diversity, 103–127. In Arratia, G., Schultze, H.-P. & Wilson, M.V.H. (eds) Mesozoic fishes 4. Homology and Phylogeny. Dr. Friedrich Pfeil, Munchen.

Mutter, R.J. & Rieber, H. 2005. Pyknotylacanthus spathianus gen. et sp. nov., a new ctenacanthoid from the Early Triassic of Bear Lake (Idaho, USA). Revista Brasileira de Paleontologia 8, 139–148. View article

Nawrocki, J. & Szulc, J. 2000. The Middle Triassic magnetostratigraphy from the Peri-Tethys basin in Poland. Earth and Planetary Science Letters 182(1), 77–92. View article

Neuman, A.G. & Wilson, M.V.H. 1985. A fossil fish of the family Saurichthyidae from the Lower Jurassic of western Alberta, Canada. Canadian Journal of Earth Sciences 22, 1158–1162. View article

Newell, N.D. & Kummel, B. 1942. Lower Eo-Triassic stratigraphy, western Wyoming and southeast Idaho. Bulletin of the Geological Society of America 53, 937–996.

Nielsen, E. 1936. Some few preliminary remarks on Triassic Fishes from East Greenland. Meddelelser om Gronland 112(3), 1–55.

Novikov, I.V., Sennikov, A.G., Minikh, A.V., Minikh, M.G. & Bukina, T.F. 2002. New data on the Early Triassic vertebrate locality Donskaya Luka in the Volgograd region. Paper II [Novye dannye po mestonahozhdeniyu rannetriasovyh poswonochnyh Donskaya Luka (Volgogradskaya oblast’). Statya II]. Geologija i razvedka 2002(2), 43–53. [in Russian, with English abstract]

Oertle, G.F. 1928. Das Vorkommen von Fischen in der Trias Wurttembergs. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie B 60, 325–472.

Olsen, P.E., McCune, A.R. & Thomson, K.S. 1982. Correlation of the Early Mesozoic Newark Supergroup by vertebrates, principally fishes. American Journal of Science 282, 1–44. View article

Oosterink, H.W. 1986. Winterswijk, geologie 2. De trias-periode (geologie, mineralen en fossielen). Wetenschappelijke Mededelingen van de Koninklijke Nederlandse Natuurhistorische Vereniging 178, 1–120.

Orchard, M.J. 2007. Conodont diversity and evolution through the latest Permian and Early Triassic upheavals. Palaeogeography, Palaeoclimatology, Palaeoecology 252, 93–117. View article

Orvig, T. 1978. Microstructure and growth of the dermal skeleton in fossil actinopterygian fishes: Birgeria and Scanilepis. Zoologica Scripta 7, 33–56. View article

Ovtcharova, M., Bucher, H., Schaltegger, H., Galfetti, T. Brayard, A. & Guex, J. 2006. New Early to Middle Triassic U-Pb ages from south China: calibration with ammonoid biochronozones and implications for the timing of the Triassic biotic recovery. Earth and Planetary Science Letters 243, 463–475. View article

Owen, R. 1860. Palaeontology or A systematic summary of extinct animals and their geological relations. 420 pp. A. and C. Black, Edinburgh. View article

Patterson, C. 1982. Morphology and interrelationships of primitive actinopterygian fishes. American Zoologist 22(2), 241–259.

Patton, W.W. Jr. & Tailleur, I.L. 1964. Geology of the Killik-Itkillik region, Alaska 3, areal geology. US Geological Survey, Professional Paper 303-G, 409–500.

Paull, R.A. & Paull, R.K. 1993. Interpretation of Early Triassic nonmarine-marine relations, Utah, U.S.A., 403–409. In Lucas, S.G. & Morales,M. (eds) The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin 3.

Piveteau, J. 1944–1945. Paleontologie de Madagascar XXV – Les poissons du Trias inferieur, la famille des Saurichthyides. Annales de Paléontologie 31, 79–89.

Preto, N., Kustatscher, E. & Wignall, P.B. 2010. Triassic climates – State of the art and perspectives. Palaeogeography, Palaeoclimatology, Palaeoecology 290, 1–10. View article

Raymond, P.E. 1925. Two new fossil fishes from Alberta. The American Journal of Science 5(9), 551–555. View article

Reeside, J.B. Jr., Applin, P.L., Colbert, E.H., Gregory, J.T., Hadley, H.D., Kummel, B., Lewis, P.J., Love, J.D., Maldonado-Koerdell, M., McKee, E.D., McLaughlin, D.B., Muller, S.W., Reinemund, J.A., Rodgers, J., Sanders, J., Silberling, N.J. & Waage, K. 1957. Correlation of the Triassic formations of North America exclusive Canada. Bulletin of the Geological Society of America 68, 1451–1524. View article

Reis, O.M. 1892. Zur Osteologie und Systematik der Belonorhynchiden und Tetragonolepiden. Geognostische Jahreshefte 4, 143–171.

Renesto, S. & Stockar, R. 2009. Exceptional preservation of embryos in the actinopterygian Saurichthys from the Middle Triassic of Monte San Giorgio, Switzerland. Swiss Journal of Geoscience 102, 323–330. View article

Rieppel, O. 1980a. Additional specimens of Saurichthys madagascariensis Piveteau, from the Eotrias of Madagascar. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1980(1), 43–51.

Rieppel, O. 1980b. A new coelacanth from the Middle Triassic of Monte San Giorgio, Switzerland. Eclogae Geologicae Helvetiae 73(3), 921–939.

Rieppel, O. 1985. Die Triasfauna der Tessiner Kalkalpen XXV. Die Gattung Saurichthys (Pisces, Actinopterygii) aus der mittleren Trias des Monte San Giorgio, Kanton Tessin. Schweizerische Paläontologische Abhandlungen 108, 1–103.

Rieppel, O. 1992. A new species of the genus Saurichthys (Pisces: Actinopterygii) from the Middle Triassic of Monte San Giorgio (Switzerland), with comments on the phylogenetic interrelationships of the genus. Palaeontographica A 221, 63–94.

Rieppel, O. 2000. Middle Triassic marine vertebrates from the northern Gondwanan shelf. Zentralblatt für Geologie und Paläontologie 1 (9/10), 1269–1284.

Rieppel, O., Kindlimann, R. & Bucher, H. 1996. A new fossil fish fauna from the Middle Triassic (Anisian) of north-western Nevada, 501–512. In Arratia, G. & Viohl, G. (eds) Mesozoic Fishes, Systematics and Paleoecology. Dr. Friedrich Pfeil, Munchen.

Ritchie, A. 1981. First complete specimen of the dipnoan Gosfordia truncata Woodward from the Triassic of New South Wales. Records of the Australian Museum 33(11), 606–615. View article

Ritchie, A. 1987. The great Somersby fossil fish dig. Australian Natural History 22(4), 146–150.

Romer, A.S. 1955. Herpetichthyes, Amphibioidei, Choanichthyes or Sarcopterygii? Nature 176, 126. View article

Rosen, D.E., Forey, P.L., Gardiner, B.G. & Patterson, C. 1981. Lungfishes, tetrapods, paleontology, and plesiomorphy. Bulletin of the American Museum of Natural History 169, 159–276.

Sander, P.M., Rieppel, O.C. & Bucher, H. 1994. New marine vertebrate fauna from the Middle Triassic of Nevada. Journal of Paleontology 68(3), 676–680.

Schaeffer, B. & Mangus, M. 1976. An Early Triassic fish assemblage from British Columbia. Bulletin of the American Museum of Natural History 156, 127–216.

Schaltegger, U., Guex, J., Bartolini, A., Schoene, B. & Ovtcharova, M. 2008. Precise U-Pb age constraints for end-Triassic mass extinction, its correlation to volcanism and Hettangian post-extinction recovery. Earth and Planetary Science Letters 267, 266–275. View article

Schmid, E.E. 1846. Die geognostischen Verhaltnisse des Saalthales bei Jena, 1–72. In Schmid, E.E. & Schleiden, M.J. (eds) Die geognostischen Verhältnisse des Saalthales bei Jena. Wilhelm Engelmann, Leipzig.

Schmid, E.E. 1861. Die Fischzähne der Trias bei Jena. 42 pp. Friedrich Frommann, Jena.

Schmid, L. & Sanchez-Villagra, M.R. 2010. Potential genetic bases of morphological evolution in the Triassic fish Saurichthys. Journal of Experimental Zoology (Molecular and Developmental Evolution) 314B, 519–526. View article

Schultze, H.-P. 2004. Mesozoic sarcopterygians, 463–492. In Arratia, G. & Tintori, A. (eds) Mesozoic fishes 3. Systematics, Paleoenvironments and Biodiversity. Dr. Friedrich Pfeil, Munchen.

Schultze, H.-P. & Moller, H. 1986. Wirbeltierreste aus dem Mittleren Muschelkalk (Trias) von Gottingen, West-Deutschland. Paläontologische Zeitschrift 60(1/2), 109–129.

Seegis, D. 1997. Die Lehrbergschichten im Mittleren Keuper von Süddeutschland – Stratigraphie, Petrographie, Paläontologie, Genese. Geo-Regio Forschung 1. 382 pp. Dr. Manfred Hennecke, Remshalden.

Sennikov, A.G. 1996. Evolution of the Permian and Triassic tetrapod communities of Eastern Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 120, 331–351. View article

Silberling, N.J. & Tozer, E.T. 1968. Biostratigraphic classification of the marine Triassic in North America. Special Paper of the Geological Society of America 110, 1–63.

Stensio, E. 1919. Einige Bemerkungen uber die systematische Stellung von Saurichthys mougeoti Agassiz. Senckenbergiana 1(6), 177–181.

Stensio, E. 1921. Triassic fishes from Spitzbergen 1. 307 pp. Adolf Holzhausen, Wien.

Stensio, E. 1925. Triassic fishes from Spitzbergen 2. Kungliga Svenska Vetenskapsakademiens Handlingar 3(2), 1–261.

Stensio, E. 1932. Triassic fishes from East Greenland 1–2. Meddelelser om Gronland 83(3), 1–298.

Storrs, G.W. 1994. Fossil vertebrate faunas of the British Rhaetian (Latest Triassic). Zoological Journal of the Linnean Society 112, 217–259. View article

Sytchevskaya, E.K. 1999. Freshwater fish fauna from the Triassic of northern Asia, 445–468. In Arratia, G. & Schultze, H.-P. (eds) Mesozoic Fishes 2. Systematics and Fossil Record. Dr. Friedrich Pfeil, Munchen.

Tanner, V.M. 1936. A study of Utah fossil fishes with the description of a new genus and species. Proceedings of the Utah Academy of Sciences, Arts and Letters 13, 81–89.

Thies, D. 1985. Funde von Acidorhynchus brevirostris (Woodward 1895) aus dem Posidonienschiefer (Unter-Toarcium) NW-Deutschlands. Palaeontographica A 187, 183–203.

Tintori, A. 1990. The vertebral column of the Triassic fish Saurichthys (Actinopterygii) and its stratigraphical significance. Rivista Italiana di Paleontologia e Stratigrafia 96(1), 93–102.

Tintori, A. 1998. Fish biodiversity in the marine Norian (Late Triassic) of northern Italy: the first Neopterygian radiation. Italian Journal of Zoology 65(1), 193–198. View article

Tintori, A. & Gozzi, E. 2005. Proposal for a taxonomical revision of Norian species of the genus Saurichthys, 249–252. In Poyato-Ariza, F.J. (ed.) Fourth International Meeting on Mesozoic fishes. Systematics, Homology, and Nomenclature. Extended abstracts. Universidad Autonoma de Madrid, Madrid.

Tintori, A. & Lombardo, C. 1999. Late Ladinian fish faunas from Lombardy (North Italy): stratigraphy and paleobiology, 495–504. In Arratia, G. & Schultze, H.-P. (eds) Mesozoic Fishes 2. Systematics and Fossil Record. Dr. Friedrich Pfeil, Munchen.

Tintori, A., Muscio, G. & Nardon, S. 1985. The Triassic fossil fishes localities in Italy. Rivista Italiana di Paleontologia e Stratigrafia 91(3), 197–210.

Tong, J., Zhou, X., Erwin, D. H., Zuo, J. & Zhao, L. 2006. Fossil fishes from the Lower Triassic of Majiashan, Chaohu, Anhui Province, China. Journal of Paleontology 80(1), 146–161. View article

Turner, S. 1982. Saurichthys (Pisces, Actinopterygii) from the Early Triassic of Queensland. Memoirs of the Queensland Museum 20, 545–551.

Tverdokhlebov, V.P., Tverdokhlebova, G.I., Minikh, A.V., Surkov, M.V. & Benton, M.J. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69, 27–77. View article

Wade, R.T. 1935. The Triassic fishes of Brookvale, New South Wales. 89 pp. British Museum (Natural History), London.

Wade, R.T. 1939. The Triassic fishes of Gosford, New South Wales. Journal of the Royal Society of New South Wales 73, 206–217.

Walker, J.D. & Geissman, J.W. 2009. 2009 GSA geologic time scale. GSA Today 19(4), 60–61. View article

Wang, X., Chen, X., Wang, C. & Cheng, L. 2009. The Triassic Guanling fossil Group – A key GeoPark from a barren mountain, Guizhou Province, China, 11–28. In Lipps, J.H. & Granier, B.R.C. (eds) PaleoParks – The protection and conservation of fossil sites worldwide. Carnets de Géologie 2009(3).

Warren, A.A. 1980. Parotosuchus from the Early Triassic of Queensland and Western Australia. Alcheringa: An Australasian Journal of Palaeontology 4(1), 25–36.

Webb, P.W., Hardy, D.H. & Mehl, V.L. 1992. The effect of armored skin on the swimming of longnose gar, Lepisosteus osseus. Canadian Journal of Zoology 70(6), 1173–1179. View article

Weitschat, W. & Dagys, A.S. 1989. Triassic biostratigraphy of Svalbard and a comparison with NE-Siberia. Mitteilungen des Geologisch-Paläontologischen Instituts der Universität Hamburg 68, 179–213.

Wendruff, A.J. & Wilson, M.V.H. 2012. A fork-tailed coelacanth, Rebellatrix divaricerca, gen. et sp. nov. (Actinistia, Rebellatricidae, fam. nov.), from the Lower Triassic of Western Canada. Journal of Vertebrate Paleontology 32(3), 499–511. View article

Wilson, M.V.H. & Bruner, J.C. 2004. Mesozoic fish assemblages of North America, 575–595. In Arratia, G. & Tintori, A. (eds) Mesozoic fishes 3. Systematics, Paleoenvironments and Biodiversity. Dr. Friedrich Pfeil, Munchen.

Winkler, T.C. 1880. Description de quelques restes de poissons fossiles des terrains triasiques des environs de Wurzbourg. Archives du Musée Teyler 5, 109–149.

Woodward, A.S. 1888. Note on the early Mesozoic ganoid, Belonorhynchus, and on the supposed Liassic genus Amblyurus. Annals and Magazine of Natural History 6, 354–356. View article

Woodward, A.S. 1890. The fossil fishes of the Hawkesbury series at Gosford. Memoirs of the Geological Survey of New South Wales, Palaeontology 4, 1–55.

Woodward, A.S. 1895. Catalogue of the fossil fishes in the British Museum (Natural History) 3. 544 pp. British Museum (Natural History), London.

Woodward, A.S. 1912. Notes on some fish-remains from the Lower Trias of Spitzbergen. Bulletin of the Geological Institute of Upsala 11, 291–298.

Wu, F.X., Sun, Y.L., Hao, W.C., Jiang, D.Y., Xu, G.H., Sun, Z.Y. & Tintori, A. 2009. A new species of Saurichthys (Actinopterygii: Saurichthyidae) from Middle Triassic (Anisian) of Yunnan Province, China. Acta Geologica Sinica, English Edition 83, 440–450.

Wu, F.X., Sun, Y.L., Xu, G.H., Hao, W.C., Jiang, D.Y. & Sun, Z.Y. 2011. New saurichthyid actinopterygian fishes from the Anisian (Middle Triassic) of southwestern China. Acta Palaeontologica Polonica 56(3), 581–614. View article

Youngquist, W. 1952. Triassic conodonts from southeastern Idaho. Journal of Paleontology 26(4), 650–655.

Zhang, Q.Y., Zhou, C.Y., Lu, T. & Bai, J.K. 2010. Discovery of Middle Triassic Saurichthys in the Luoping area, Yunnan, China. Geological Bulletin of China 29, 26–30. [in Chinese, with English abstract]

Zittel, K.A. von 1887–1890. Handbuch der Palaeontologie. 1. Abtheilung. Palaeozoologie. III. Vertebrata. 900 pp. R. Oldenbourg, Munchen & Leipzig.