A new selenosteid arthrodire (‘Placodermi’) from the Late Devonian of Morocco

LITERATURE CITED

• Ahlberg, P. E., K. Trinajstic, Z. Johanson, and J. A. Long. 2009. Pelvic claspers confirm chondrichthyan-like internal fertilization in arthrodires. Nature 460:888–89.
   PubMed Web of Science ®Google Scholar
• Asmuss, H. 1856. Das vollkommenste Hautskelett der bisher bekannten Thierreihe, An fossilen Fischen des Alten Rothen Sandsteins aufgefunden und aus ihren Resten erläutert. Abhandlung zur Erlangung der Magisterwürd. Gedruckt bei Schünmans Wittwe and C. Mattiesen, Dorpat, 39 pp.
   Google Scholar
• Brazeau, M. D. 2009. The braincase and jaws of a Devonian ‘acanthodian’ and modern gnathostome origins. Nature 457:305–308.
   PubMed Web of Science ®Google Scholar
• Carr, R. K. 1991. Reanalysis of Heintzichthys gouldii (Newberry), an aspinothoracid arthrodire (Placodermi) from the Famennian of northern Ohio, with a review of brachythoracid systematics. Zoological Journal of the Linnean Society 103:349–390.
   Web of Science ®Google Scholar
• Carr, R. K. 1994. A redescription of Gymnotrachelus (Placodermi: Arthrodira) from the Cleveland Shale (Famennian) of Northern Ohio, USA. Kirtlandia 48:3–21.
   Google Scholar
• Carr, R. K. 1996. Stenosteus angustopectus sp. nov. from the Cleveland Shale (Famennian) of Northern Ohio with a review of selenosteid (Placodermi) systematics. Kirtlandia 49:19–43.
   Google Scholar
• Carr, R. K., and W. J. Hlavin. 2010. Two new species of Dunkleosteus Lehman, 1956, from the Ohio Shale Formation (USA, Famennian) and the Kettle Point Formation (Canada, Late Devonian) and a cladistic analysis of the Eubrachythoraci (Placodermi, Arthrodira). Zoological Journal of the Linnean Society 159:195–222.
   Web of Science ®Google Scholar
• Davis, S. P., J. A. Finarelli, and M. I. Coates. 2012. Acanthodes and shark-like conditions in the last common ancestor of modern gnathostomes. Nature 486:247–251.
   PubMed Web of Science ®Google Scholar
• Dean, B. 1901. III. Further notes on the relationships of the Arthrognathi. Memoirs of the New York Academy of Sciences 2:110–123.
   Google Scholar
• Denison, R. H. 1978. Placodermi. Handbook of Paleoichthyology, Volume 2. Gustav Fischer Verlag, Stuttgart, 128 pp.
   Google Scholar
• Dennis-Bryan, K. 1987. A new species of eastmanosteid arthrodire (Pisces: Placodermi) from Gogo, Western Australia. Zoological Journal of the Linnean Society 90:1–64.
   Web of Science ®Google Scholar
• Dunkle, D. H, and P. A. Bungart. 1939. A new arthrodire from the Cleveland Shale Formation. Scientific Publications of the Cleveland Museum of Natural History 8:13–28.
   Google Scholar
• Feist, R. 2002. Trilobites from the latest Frasnian Kellwasser Crisis in North Africa (Mrirt, central Moroccan Meseta). Acta Palaeontologica Polonica 47:203–210.
   Web of Science ®Google Scholar
• Gardiner, B. G., and R. S. Miles. 1994. Eubrachythoracid arthrodires from Gogo, Western Australia. Zoological Journal of the Linnean Society 112:443–477.
   Web of Science ®Google Scholar
• Goujet, D. F. 1984. Les Poissons Placodermes du Spitsberg, Arthrodires Dolichothoraci de la Formation de Wood Bay (Dévonien Inférieur). Centre National de la Recherche Scientifique, Paris, 284 pp.
   Google Scholar
• Goujet, D. 2001. Placoderms and basal gnathostome apomorphies; pp. 209–222 in P. E. Ahlberg (ed.), Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny Genetics and Development. Systematic Association Special Volume Series, 61. Taylor and Francis, London.
   Google Scholar
• Goujet, D., and G. C. Young. 2004. Placoderm anatomy and phylogeny: new insights; pp. 109–126 in G. Arratia, M. V. H. Wilson, and R. Cloutier (eds.), Recent Advances in the Origin and Early Radiation of Vertebrates. Verlag Dr. Friedrich Pfeil, Munich.
   Google Scholar
• Gross, W. 1932. Die Arthrodira Wildungens. Geologische und Paläontologische Abhandlungen, Neue Folge, Jena 19:1–61.
   Google Scholar
• Hairapetian, V., J. Valiukevicius, and C. J. Burrow. 2006. Early Frasnian acanthodians from central Iran. Acta Palaeontologica Polonica 51:499–520.
   Web of Science ®Google Scholar
• Heintz, A. 1932. The structure of Dinichthys: a contribution to our knowledge of the Arthrodira. Bashford Dean Memorial Volume—Archaic Fishes 4:115–224.
   Google Scholar
• Jaekel, O. M. J. 1903. Über die Organisation und systematische Stellung der Asterolepiden. Zeitschrift der Deutschen Geologischen Gesellschaft 55:41–60.
   Google Scholar
• Jaekel, O. M. J. 1911. Die Wirbeltiere. Gebrüder Bornträger, Berlin, 252 pp.
   Google Scholar
• Jaekel, O. M. J. 1919. Die Mundbildung der Placodermen. Sitzungsberichte der Gesellschaft Naturforschender Freunde, Berlin 1919:73–110.
   Google Scholar
• Janvier, P., and C. Villarroel. 2000. Devonian vertebrates from Colombia. Palaeontology 43:729–763.
   Web of Science ®Google Scholar
• Janvier, P., and J. G. Maisey. 2010. The Devonian vertebrates of South America and their biogeographical relationships; pp. 433–461 in D. K. Elliot, J. G. Maisey, X. Yu, and D. Miao (eds.), Morphology, Phylogeny and Paleobiogeography of Fossil Fishes. Verlag Dr. Friedrich Pfeil, Munich.
   Google Scholar
• Klug, C., B. Kröger, M. Rücklin, D. Korn, M. Schemm-Gregory, K. De Baets, and R. H. Mapes. 2008. Ecological change during the early Emsian (Devonian) in the Tafilalt (Morocco), the origin of the Ammonoidea, and the first African pyrgocystid edrioasteroids, machaerids and phyllocarids. Palaeontographica A 283:1–94.
   Google Scholar
• Lehman, J. P. 1956. Les arthrodires du Dévonien supérieur du Tafilalt (Sud marocain). Notes et Memoires du Service Géologique du Maroc 129:1–70.
   Google Scholar
• Lehman, J. P. 1976. Nouveaux poissons fossiles du Dévonien du Maroc. Annales de Paléontologie, Vertébrés 62:1–34.
   Google Scholar
• Lehman, J. P. 1977. Nouveaux arthrodires du Tafilalt et de ses environs. Annales de Paléontologie, Vertébrés 63:105–132.
   Google Scholar
• Lelièvre, H. 1995. Description of Maideria falipoui n. g., n. sp., a long snouted brachythoracid (Vertebrata, Placodermi, Arthrodira) from the Givetian of Maider (South Morocco), with a phylogenetic analysis of primitive brachythoracids. Bulletin du Muséum national d’histoire naturelle, Paris, 4ème série, Section C 17:163–207.
   Google Scholar
• Lelièvre, H., P. Janvier, and A. Blieck. 1993. Silurian-Devonian vertebrate biostratigraphy of western Gondwana and related terranes (South America, Africa, Armorica-Bohemia, Middle East); pp. 139–173 in J. A. Long (ed.), Palaeozoic Vertebrate Biostratigraphy and Biogeography. Belhaven Press, London.
   Google Scholar
• Lelièvre, H., R. Feist, D. Goujet, and A. Blieck. 1987. Les vertébrés de la Montagne Noire (Sud de la France) et leur apport à la phylogénie des Pachyosteomorphes (Placodermes, Arthrodires). Palaeovertebrata 17:1–26.
   Google Scholar
• Long, J. A. 1988. A new camuropiscid arthrodire (Pisces, Placodermi) from Gogo, Western Australia. Zoological Journal of the Linnean Society of London 94:233–258.
   Web of Science ®Google Scholar
• Long, J. A. 1990. Two New Arthrodires (Placoderm Fishes) from the Upper Devonian Gogo Formation, Western Australia. Memoirs of the Queensland Museum 28, De Vis Symposium Volume, pp. 51–64.
   Google Scholar
• Long, J. A. 1994. A second incisoscutid arthrodire (Pisces, Placodermi) from the Late Devonian Gogo Formation, Western Australia. Alcheringa 18:59–69.
   Web of Science ®Google Scholar
• Long, J. A. 1995. A plourdosteid arthrodire from the Upper Devonian Gogo Formation of Western Australia. Palaeontology 38:39–62.
   Web of Science ®Google Scholar
• Long, J. A., and K. Trinajstic. 2010. The Late Devonian Gogo Formation Lagerstätte of Western Australia: exceptional early vertebrate preservation and diversity. Annual Review of Earth and Planetary Sciences 38:255–279.
   Web of Science ®Google Scholar
• Long, J. A., K. Trinajstic, and Z. Johanson. 2009. Devonian arthrodire embryos and the origin of internal fertilization in vertebrates. Nature 457:1124–1126.
   PubMed Web of Science ®Google Scholar
• Long, J. A., K. Trinajstic, G. C. Young, and T. Senden. 2008. Live birth in the Devonian Period. Nature 453:650–652.
   PubMed Web of Science ®Google Scholar
• Lubeseder, S., J. Rath, M. Rücklin, and R. Messbacher. 2010. Controls on Devonian hemi-pelagic limestone deposition analyzed on cephalopod ridge to slope sections, Eastern Anti-Atlas, Morocco. Facies 56:295–315.
   Web of Science ®Google Scholar
• Maisch, M. W. 1998. Wildungenichthys grossi n. gen., n. sp.—a new selenosteid arthrodire (Placodermi, Arthrodira) from the Kellwasserkalk (late Frasnian, Upper Devonian) of Bad Wildungen (Hessen, W-Germany). Paläontologische Zeitschrift 72:373–382.
   Google Scholar
• McCoy, F. 1848. On some new fossil fish of the Carboniferous Period. Annals and Magazine of Natural History 2:115–134.
   Google Scholar
• McKerrow, W. S., C. Mac Niocaill, P. E. Ahlberg, G. Clayton, C. J. Cleal, and R. M. C. Eagar. 2000. The late Palaeozoic relations between Gondwana and Laurussia; pp. 9–20 in W. Franke, V. Haak, O. Oncken, and D. Tanner (eds.), Orogenic Processes: Quantification and Modeling in the Variscan Belt. Geological Society, London, Special Publications 179.
   Google Scholar
• Meyer-Berthaud, B., J. Wendt, and J. Galtier. 1997. First record of a large Callixylon trunk from the Late Devonian of Gondwana. Geological Magazine 134:847–853.
   Web of Science ®Google Scholar
• Meyer-Berthaud, B., M. Rücklin, A. Soria, Z. Belka, and H. Lardeux. 2004. Frasnian plants from the Dra Valley, southern Anti-Atlas, Morocco. Geological Magazine 141:675–686.
   Web of Science ®Google Scholar
• Miles, R. S. 1971. The Holonematidae (placoderm fishes), a review based on new specimens of Holonema from the Upper Devonian of Western Australia. Philosophical Transactions of the Royal Society B 263:101–234.
   Web of Science ®Google Scholar
• Miles, R. S., and K. D. Dennis. 1979. A primitive eubrachythoracid arthrodire from Gogo, Western Australia. Zoological Journal of the Linnean Society 66:31–62.
   Web of Science ®Google Scholar
• Newberry, J. S. 1885. Descriptions of some gigantic placoderm fishes recently discovered in the Devonian of Ohio. Transactions of the New York Academy of Sciences 5:25–28.
   Google Scholar
• Rixon, A. E. 1976. Fossil Animal Remains: Their Preparation and Conservation. Athlone Press, London, 304 pp.
   Google Scholar
• Rücklin, M. 2010. A new Frasnian placoderm assemblage from the eastern Anti-Atlas, Morocco, and its palaeobiogeographical implications. Palaeoworld 19:87–93.
   Google Scholar
• Rücklin, M. 2011. First selenosteid placoderms from the eastern Anti-Atlas of Morocco: osteoleogy, phylogeny and palaeoegographical implications. Palaeontology 54:25–62.
   Web of Science ®Google Scholar
• Schultze, H.-P., and R. Cloutier. 1996. Comparison of the Escuminac Formation ichthyofauna with other late Givetian/early Frasnian ichthyofaunas; pp. 348–374 in H.-P. Schultze and R. Cloutier (eds.), Devonian Fishes and Plants of Miguasha, Quebec, Canada. Verlag Dr. Friedrich Pfeil, Munich.
   Google Scholar
• Stensiö, E. A. 1944. Contributions to the knowledge of the vertebrate fauna of the Silurian and Devonian of western Podolia. II. Notes on two arthrodires from the Downtonian of Podolia. Arkiv foer Zoologi 35:1–83.
   Google Scholar
• Stensiö, E. A. 1959. On the pectoral fin and shoulder girdle of the arthrodires. Kungliga Svenska Vetenskapsakademiens Handlingar 8:1–229.
   Google Scholar
• Stensiö, E. A. 1963. Anatomical studies on the arthrodiran head. Part 1. Kungliga Svenska Vetenskapsakademiens Handlingar, Series 4 9:1–419.
   Google Scholar
• Swofford, D. A. 2003. PAUP* 4.0. Sinauer Associates, Sunderland, Massachusetts.
   Google Scholar
• Toombs, H. A., and A. E. Rixon. 1959. The use of acids in the preparation of vertebrate fossils. Curator: The Museum Journal 2:304–312.
   Google Scholar
• Trinajstic, K., and K. Dennis-Bryan. 2009. Phenotypic plasticity, polymorphism and phylogeny within placoderms. Acta Zoologica 90:83–102.
   Web of Science ®Google Scholar
• Trinajstic, K., and A. George. 2009. Microvertebrate biostratigraphy of Upper Devonian (Frasnian) carbonate rocks in the Canning and Carnarvon basins of Western Australia. Palaeontology 52:641–659.
   Web of Science ®Google Scholar
• Trinajstic, K. M., and M. Hazelton. 2007. Ontogeny, phenotypic variation and phylogenetic implications of arthrodires from the Gogo Formation, Western Australia. Journal of Paleontology 27:571–583.
   Web of Science ®Google Scholar
• Trinajstic, K., and J. A. Long. 2009. A new genus and species of ptyctodont (Placodermi) from the Late Devonian Gneudna Formation, Western Australia, and an analysis of ptyctodont phylogeny. Geological Magazine 146:743–760
   Web of Science ®Google Scholar
• Trinajstic, K., J. A. Long, Z. Johanson, G. Young and T. Senden. 2012. New morphological information on the ptyctodontid fishes (Placodermi, Ptyctodontida) from Western Australia. Journal of Vertebrate Paleontology 32:757–780
   Web of Science ®Google Scholar
• Turner, S., C. Burrow, H. Gholamalian, and M. Yazdi. 2002. Late Devonian (early Frasnian) microvertebrates and conodonts from the Chahriseh area near Esfahan, Iran. Memoirs of the Association of Australasian Paleontologists 27:149–159.
   Google Scholar
• Wang, C.-C. 1979. A new family of Arthrodira from Yunnan, China. Vertebrata PalAsiatica 17:179–188.
   Web of Science ®Google Scholar
• Wendt, J., and Z. Belka. 1991. Age and Depositional Environment of Upper Devonian (early Frasnian to early Famennian) Black Shales and Limestones (Kellwasser Facies) in the Eastern Anti-Atlas, Morocco. Facies 25:51–90.
   Google Scholar
• Woodward, A. S. 1891. Catalogue of the Fossil Fishes in the British Museum (Natural History) Part II. British Museum (Natural History), London, 567 pp.
   Google Scholar
• Young, G. C. 1979. New information on the structure and relationships of Buchanosteus (Placodermi: Euarthrodira) from the Early Devonian of New South Wales. Zoological Journal of the Linnean Society of London 66:309–352.
   Web of Science ®Google Scholar
• Young, G. C. 1981. New Early Devonian brachythoracids (placoderm fishes) from the Taemas–Wee Jasper region of New South Wales. Alcheringa 5:245–271.
   Web of Science ®Google Scholar
• Young, G. C. 1987. Devonian palaeontological data and the Armorica problem. Palaeogeography, Palaeoclimatology, Palaeoecology 60:283–304.
   Web of Science ®Google Scholar
• Young, G. C. 1988. Antiarchs (placoderm fishes) from the Devonian Aztec Siltstone, southern Victoria Land, Antarctica. Palaeontographica Abteilung A 202:1–125
   Google Scholar
• Young, G. C. 2003a. North Gondwana mid-Palaeozoic connections with Euramerica and Asia: Devonian vertebrate evidence. Courier Forschungsinstitut Senckenberg 242:169–185.
   Google Scholar
• Young, G. C. 2003b. A new species of Atlantidosteus Lelièvre, 1984 (Placodermi, Arthrodira, Brachythoraci) from the Middle Devonian of the Broken River area (Queensland, Australia). Geodiversitas 25:681–694.
   Web of Science ®Google Scholar
• Young, G. C. 2005. An articulated phyllolepid fish (Placodermi) from the Devonian of central Australia: implications for non-marine connections with the Old Red Sandstone continent. Geological Magazine 142:173–186.
   Web of Science ®Google Scholar
• Young, G. C. 2010. Placoderms (armored fish): dominant vertebrates of the Devonian period. Annual Review of Earth and Planetary Sciences 38:523–550.
   Web of Science ®Google Scholar
• Young, G. C., and J. Mody. 2002. A Middle-Late Devonian fish fauna from the Sierra de Perijá, western Venezuela, South America. Mitteilungen aus dem Museum für Naturkunde in Berlin. Geowissenschaftliche Reihe 5:155–206.
   Google Scholar
• Young, G. C., J. Mody, and J. Casas. 2000. New discoveries of Devonian vertebrates from South America, and implications for Gondwana-Euramerica contact. Academie des Sciences Comptes Rendus Geoscience 331:755–761.
   Web of Science ®Google Scholar
• Zhu, M., X. Yu, P. E. Ahlberg, B. Choo, J. Lu, T. Qiao, Q. Qu, W. Zhao, H. Bloom, and Y. Zhu. 2013. A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature 502:188–193.
   PubMed Web of Science ®Google Scholar
• Submitted April 16, 2013; revisions received March 11, 2014; accepted March 20, 2014.
   Google Scholar
• Handling editor: Martha Richter.
   Google Scholar