Late Pennsylvanian fish assemblage from the Robledo Mountains and new records of Paleozoic chondrichthyans in New Mexico, USA


Authors: Ivanov AO, Lucas SG

Published in: Bulletin of Geosciences, volume 94, issue 2; pages: 235 - 255; Received 1 January 2019; Accepted in revised form 4 April 2019; Online 10 May 2019

Keywords: fishes, Late Devonian-early Permian, New Mexico,

full text (PDF, 7.82 MB)

Export to RIS



We describe a very diverse and rich assemblage of fossil fishes from the Upper Pennsylvanian (Missourian) interval of the Horquilla Formation in the Robledo Mountains of southern New Mexico, USA. The assemblage includes bransonelliforms, symmoriiforms, a ctenacanthiform, a jalodontid, euselachians, neoselachians, an orodontiform, a helodontiform, an eugeneodontiform, a petalodontiform, a psephodontid, an acanthodian and actinopterygians. For the first time, the vascularization system of the teeth of Adamantina, Bransonella, Helodus and Agassizodus has been studied using a micro-CT. Besides the diverse fish assemblage from the Robledo Mountains, some other new records of Paleozoic chondrichthyans in New Mexico are documented.


Agassiz, L. 1833-43. Recherches sur les Poissons fossils. 5 vols + supplement. 1420 pp. Petitpierre, Neuchâtel.

Barrick, J.E., Lambert, L.L., Heckel, P.H., Rosscoe, S.J. & Boardman, D.R. 2013. Midcontinent Pennsylvanian conodont zonation. Stratigraphy 10, 55-72.

Behan, C., Walken, G. & Cuny, G. 2012. A Carboniferous chondrichthyan assemblage from residues within a Triassic karst system at Cromhall Quarry, Gloucestershire, England. Palaeontology 55(6), 1245-1263.View article

Bendix-Almgreen, S.E. 1993. Adamantina benedictae n. g. et sp. - en nyhed fra Ostgronlands marine Ovre Perm, 48-58. In Johnsen, O. (ed.) Geologisk Museum - 100 ar pa Ostervold. Rodos, Kobenhavn.

Binney, E.W. 1840. On the fossil fishes of the Pendleton Coal Field. Transactions of the Geological Society of Manchester 1840, 153-178.

Bonaparte, C.L.J. 1832-1841. Iconografia della Fauna Italica, per le quattro classi degli animali vertebrati. Tomo 3: Pesci. 266 pp. Dalla Tipografia Salviucci, Rome.

Bonaparte, C.L.J. 1838. Selachorum tabula analytica. Nuovi Annali delle Scienze Naturali Bologna 1, 195-214.

Burrow, C.J., Long, J.A. & Trinajstic, K. 2009. Disarticulated acanthodian and chondrichthyan remains from the upper Middle Devonian Aztec Siltstone, southern Victoria Land, Antarctica. Antarctic Science 21, 71-88.View article

Cappetta, H., Duffin, C. & Zidek, J. 1993. Chondrichthyes, 593-609. In Benton, M.J. (ed.) The Fossil Record 2. Chapman and Hall, London.

Coates, M.I. & Sequeira, S.E.K. 2001. A new stethacanthid chondrichthyan from the Lower Carboniferous of Bearsden, Scotland. Journal of Vertebrate Paleontology 21, 438-459.View article

Compagno, L.J.V. 1977. Phyletic relationships of living sharks and rays. American Zoologist 17, 303-322.View article

Cope, E.D. 1884. On the structure of the skull in the elasmobranch genus Didymodus. Proceedings of the American Philosophical Society 21, 572-590.

Cope, E.D. 1888. Systematic catalogue of the species of Vertebrata found in the beds of the Permian epoch in North America, with notes and descriptions. Transactions of the American Philosophical Society 26, 285-297.View article

Cuny, G. & Stemmerik, L. 2018. New fossil fish microremains from the Upper Carboniferous of eastern North Greenland. Bulletin of the Geological Society of Denmark 66, 47-60.View article

Dean, B. 1909. Studies on fossil fishes (sharks, chimaeroids and arthrodires). Memoirs of the American Museum of Natural History Part V, 9, 211-287.

Dick, J.R.F. 1998. Sphenacanthus, a Palaeozoic freshwater shark, 9-25. In Norman, D.B., Milner, A.R. & Milner, A.C. (eds) A Study of Fossil Vertebrates. Zoological Journal of the Linnean Society 122.

Duffin, C.J. & Ward, D.J. 1983. Neoselachian shark teeth from the Lower Carboniferous of Britain and the Lower Permian of the USA. Palaeontology 26(1), 93-110.

Duffin, C.J., Richter, M. & Neis, P.A. 1996. Shark remains from the Late Carboniferous of the Amazon Basin, Brazil. Neues Jahrbuch für Geologie und Paläontologie 4, 232-256.View article

Duncan, M. 2004. Chondrichthyan genus Lissodus from the Lower Carboniferous of Ireland. Acta Palaeontologica Polonica 49(3), 417-428.

Eastman, C.R. 1902. Some Carboniferous cestraciont and acanthodian sharks. Bulletin of the Museum of Comparative Zoology 39(3), 55-99.

Giebel, C.G. 1848. Fauna der Vorwelt mit steter Berücksichtigung der lebenden Thiere, Bd. 1: Wirbelthiere, Abt. 3: Fische. 467 pp. Brockhaus, Leipzig.

Ginter, M. 1990. Late Famennian shark teeth from the Holy Cross Mts, Central Poland. Acta Geologica Polonica 40, 69-81.

Ginter, M. & Hansen, M. 2010. Teeth of the cladodont shark Denaea from the Carboniferous of central North America, 29-44. In: Nowakowski, D. (ed.) Morphology and Systematics of Fossil Vertebrates. DN Publisher, Wroclaw.

Ginter, M. & Ivanov, A. 1995. Middle/Late Devonian phoebodont-based ichthyolith zonation. Géobios, Mémoire Special 19, 351-355.View article

Ginter, M., Hairapetian, V. & Klug, C. 2002. Famennian chondrichthyans from the shelves of North Gondwana. Acta Geologica Polonica 52, 169-215.

Ginter, M., Hampe, O. & Duffin, C.J. 2010. Chondrichthyes Paleozoic Elasmobranchii: Teeth, 1-168. In Schultze, H.-P. (ed.) Handbook of Paleoichthyology, 3D. Verlag Dr. Friedrich Pfeil, Munich.

Ginter, M., Ivanov, A. & Lebedev, O. 2005. The revision of “Cladodusoccidentalis, a late Palaeozoic ctenacanthiform shark. Acta Palaeontologica Polonica 50, 623-631.

Ginter, M., Duffin, C.J., Dean, M.T. & Korn, D. 2015. Late Viséan pelagic chondrichthyans from northern Europe. Acta Palaeontologica Polonica 60(4), 899-922.

Glikman, L.S. 1964. Subclass Elasmobranchii. Sharks, 196-237. In Obruchev, D.V. (ed.) Fundamentals of Paleontology. Agnathans, Fishes. Nauka, Moscow. [in Russian]

Grogan, E.D. & Lund, R. 2000. Debeerius ellefseni (Fam. Nov., Gen. Nov., Spec. Nov.), an autodiastylic chondrichthyan from the Mississippian Bear Gulch Limestone of Montana (USA), the relationships of the Chondrichthyes, and comments on gnathostome evolution. Journal of Morphology 243, 219-245.View article

Gross, W. 1938. Das Kopfskelett von Cladodus wildungensis Jaekel. 2. Teil: Der Kieferbogen. Anhang: Protacrodus vetustus Jaekel. Senckenbergiana 20, 123-145.

Gunnell, F.M. 1933. Conodonts and fish remains from the Cherokee, Kansas City, and Wabaunsee groups of Missouri and Kansas. Journal of Paleontology 7, 261-297.

Habibi, T. & Ginter, M. 2011. Early Carboniferous chondrichthyans from the Mobarak Formation, central Alborz Mountains, Iran. Acta Geologica Polonica 61, 27-34.

Hampe, O. 2003. Revision of the Xenacanthida (Chondrichthyes: Elasmobranchii) from the Carboniferous of the British Isles. Transactions of the Royal Society of Edinburgh: Earth Sciences 93(3), 191-237.View article

Hampe, O. & Ivanov, A. 2007. Bransonelliformes - a new order of the Xenacanthimorpha (Chondrichthyes: Elasmobranchii). Fossil Records 10(2), 190-194.View article

Hampe, O., Hairapetian, V., Dorka, M., Witzmann, F., Akbari, A.M. & Korn, D. 2013. A first Late Permian fish fauna from Baghuk Mountain (Neo-Tethyan shelf, central Iran). Bulletin of Geosciences 88(1), 1-20.

Harlton, B.H. 1933. Micropaleontology of the Pennsylvanian Johns Valley shale of the Ouachita Mountains, Oklahoma, and its relationship to the Mississippian Caney shale. Journal of Paleontology 7(1), 3-29.

Hay, O.P. 1902. Bibliography and catalogue of the fossil Vertebrata of North America. U.S. Geological Survey Bulletin 179, 1-868.

Hay, O.P. 1929. Second bibliography and catalogue of the fossil Vertebrata of North America. Carnegie Institution of Washington Publication 390(1), 1-916.

Hodnett, J.-P.M. & Elliott, D.K. 2018. Carboniferous chondrichthyan assemblages from the Surprise Canyon and Watahomigi formations (latest Mississippian-Early Pennsylvanian) of the western Grand Canyon, Northern Arizona. Journal of Paleontology 92, Memoir 77, 1-33.View article

Hodnett, J-P.M. & Lucas, S.G. 2015. Paleozoic fishes of New Mexico: A review. New Mexico Museum of Natural History and Science, Bulletin 68, 51-63.

Hodnett, J.-P.M. & Lucas, S.G. 2017. Paleoichthyological assemblages of the Upper Carboniferous-Lower Permian of Socorro County, New Mexico. New Mexico Museum of Natural History and Science, Bulletin 77, 133-138.

Hodnett, J-P.M., Elliott, D.K., Olson, T.J. & Wittke, J.H. 2012. Ctenacanthiform sharks from the Permian Kaibab Formation, northern Arizona. Historical Biology 24, 381-395.View article

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 Zoological Society of London 1880, 649-662.

Itano, W.M. & Lucas, S.G. 2018. A revision of Campyloprion Eastman, 1902 (Chondrichthyes, Helicoprionidae), including new occurrences from the Upper Pennsylvanian of New Mexico and Texas, USA. Acta Geologica Polonica 68(3), 403-419.

Ivanov, A. 1999. Late Devonian - Early Permian chondrichthyans of the Russian Arctic. Acta Geologica Polonica 49, 267-285.

Ivanov, A. 2011. Permian anachronistid sharks of the East European Platform and Urals, 17-19. In Shishkin, M.A., Golubev, V.K., Novikov, I.V. & Sennikov, A.G. (eds) Palaeozoic and Mesozoic Vertebrates of Eurasia: Evolution, Assemblage Changes, Taphonomy and Palaeobiogeography. Proceedings of the International Conference. Palaeontological Institute Publ., Moscow. [in Russian]

Ivanov, A.O. 2016. Chondrichthyans from the Lower Permian of Mechetlino, South Urals, Russia. The Bulletin of Geosciences 92(4), 717-729.

Ivanov, A. & Derycke, C. 2005. Viséan elasmobranchs of Belgium. Ichthyolith Issues, Special Publication 9, 13-17.

Ivanov, A. & Ginter, M. 1996. Early Carboniferous xenacanthids (chondrichthyes) from eastern Europe. Bulletin de la Société géologique de France 167, 651-656.

Ivanov, A.O. & Hampe, O. 2015. Occurrence of xenacanthimorph sharks in the Carboniferous of Russia, 86. In Nurgaliev, D.K. (ed.) Abstract volume of the XVIII International Congress on the Carboniferous and Permian. Kazan Federal University, Kazan.

Ivanov, A.O. & Lebedev, O.A. 2014. Permian chondrichthyans of the Kanin Peninsula, Russia. Paleontological Journal 48(9), 1030-1043.View article

Ivanov, A. & Lucas, S.G. 2011. Fish fossils from the Paleozoic Sly Gap Formation of southern New Mexico, USA. New Mexico Museum of Natural History and Science, Bulletin 53, 52-70.

Ivanov, A.O. & Lucas, S.G. 2014. Fish assemblage from the late pennsylvanian of the robledo mountains, new mexico. Geological Society of America, Abstracts with Programs 46(1), 2.

Ivanov, A.O. & Nilov, S.P. 2017. Investigation of internal structure in the scales of Palaeozoic cartilaginous fishes using micro-CT. Abstracts of Micro-CT User Meeting, Brussels, 31-36.

Ivanov, A., Lucas, S.G. & Krainer, K. 2009. Pennsylvanian fishes from the Sandia Formation, Socorro County, New Mexico, 243-248. In Lueth, V.W., Lucas, S.G. & Chamberlin, R.M. (eds) Geology of the Chupadera Mesa, Guidebook 60. New Mexico Geological Society, Socorro.

Ivanov, a., Nestell, M. & Nestell, G. 2012. New jalodontid chondrichthyans from the Middle Permian of West Texas, USA. Historical Biology 24(4), 359-368.View article

Ivanov, A.O., Seuss, B. & Nützel, A. 2017. The fish assemblage from the Pennsylvanian Buckhorn Asphalt Quarry Lagerstätte (Oklahoma, USA). Paläontologische Zeitschrift 91(4), 565-576.View article

Jaekel, O. 1925. Das Mundskelett der Wirbeltiere. Gegenbaurs Morphologishes Jahrbuch 55, 402-409.

Johnson, G.D. 1981. Hybodontoidei (Chondrichthyes) from the Wichita-Albany Group (Early Permian) of Texas. Journal of Vertebrate Paleontology 1, 1-41.View article

Johnson, G.D. 1984. A new species of Xenacanthodii (Chondrichthyes, Elasmobranchii) from the late Pennsylvanian of Nebraska. Carnegie Museum of Natural History Special Publication 9, 178-186.

Johnson, G.D. 2008. Ctenacanthiform cladodont teeth from the Lower Permian Wichita Group, Texas, U.S.A. Acta Geologica Polonica 58(2), 205-209.

Johnson, G.D. 2018. Orthacanthus platypternus (Cope, 1883) (Chondrichthyes: Xenacanthiformes) teeth and other isolated vertebrate remains from a single horizon in the early Permian (Artinskian) Craddock Bonebed, lower Clear Fork Group, Baylor County, Texas, USA. Acta Geologica Polonica 68(3), 421-436.

Johnson, G.D. & Thayer, D.W. 2009. Early Pennsylvanian xenacanth chondrichthyans from the Swisshelm Mountains, Arizona, USA. Acta Palaeontologica Polonica 54(4), 649-668.View article

Kietzke, K.K. 1990. Microfossils from the Flechado Formation (Pennsylvanian, Desmoinesian) near Talpa, New Mexico, 259-276. In Bauer, P.W., Lucas, S.G., Mawer, C.K. & McIntosh, W.C. (eds) Tectonic development of the southern Sangre de Cristo Mountains, New Mexico, Guidebook 41. New Mexico Geological Society, Socorro.

Koninck, L.-G. De 1878. Faune du calcaire carbonifere de la Belgique. Premiere partie, poissons et genre nautile. Annales du Musée royal d’Histoire naturelle de Belgique 2, 1-152.

Koot, M.B., Cuny, G., Tintori, A. & Twitchett, J.J. 2013. A new diverse shark fauna from the Wordian (Middle Permian) Khuff Formation in the interior Haushi-Huqf area, Sultanate of Oman. Palaeontology 56(2), 303-343.View article

Krainer, K., Lucas, S.G., Vachard, D. & Barrick, J.E. 2015. The Pennsylvanian-Permian section at Robledo Mountain, Dona Ana County, New Mexico, USA. New Mexico Museum of Natural History and Science, Bulletin 65, 9-41.

Lebedev, O.A. 2001. Pozvonochnye [Vertebrates], 196-201. In Alekseev, A.S. & Shik, S.M. (eds) Srednij karbon Moskovskoy sineklizy (yuzhnaya chast’) 2, Paleontologicheskaya kharakteristika. Nauchniy mir, Moscow. [in Russian]

Leidy, J. 1859. Descriptions of Xystracanthus arcuatus and Cladodus occidentalis. Proceedings of the Academy of Natural Sciences of Philadelphia 1859, 3.

Lucas, S.G. & Estep, J.W. 2000. Pennsylvanian selachians from the Cerros de Amado, central New Mexico. New Mexico Museum of Natural History and Science, Bulletin 16, 21-28.

Lucas, S.G., Krainer, K. & Vachard, D. 2015. The lower Permian Hueco Group, Robledo Mountains, New Mexico. New Mexico Museum of Natural History and Science, Bulletin 65, 43-95.

Lucas, S.G., Spielmann, J.A., Ivanov, A.O., Rinehart, L.F. & Krainer, K. 2011. Petalodont chondrichthyan teeth from the Pennsylvanian-Permian Horquilla Formation, Big Hatchet Mountains, New Mexico. New Mexico Museum of Natural History and Science, Bulletin 53, 110-114.

Lund, R. & Grogan, E.D. 1997. Relationships of the Chimaeriformes and the basal radiation of the Chondrichthyes. Review Fisheries Biology Fisheries 7, 65-123.View article

Maisey, J.G. 1975. The interrelationships of phalacanthous selachians. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1975, 553-567.

Maisey, J.G. 1981. Studies on the Paleozoic selachian genus Ctenacanthus Agassiz No. 1. Historical review and revised diagnosis of Ctenacanthus, with a list of referred taxa. American Museum Novitates 2718, 1-22.

Maisey, J.G. 2010. Heslerodidae (Chondrichthyes, Elasmobranchii), a new family of Paleozoic phalacanthous sharks. Kirtlandia 37, 13-21.

Nelson, J.S. 1976. Fishes of the world. 416 pp. Wiley & Sons, New York.

Newberry, J.S. 1889. The Paleozoic fishes of North America. U.S. Geological Survey Monograph 16, 1-340.

Newberry, J.S. & Worthen, A.H. 1866. Descriptions of vertebrates. Geological Survey of Illinois 2, 9-134.

Obruchev, D.V. 1953. Edestid Studies and the Works of A.P. Karpinsky. Trudy Paleontologiceskogo Instituta Akademija Nauk SSSR 45, 1-85. [in Russian]

Ørvig, T. 1967. Histologic studies of ostracoderms, placoderms and fossil elasmobranchs. 2. On the dermal skeleton of two late Paleozoic elasmobranchs. Arkiv för Zoologi 19(1), 1-39.

Patterson, C. 1965. The phylogeny of the Chimaeroids. Philosophical Transactions Royal Society London B 249, 101-219.View article

Pruvost, P. 1922. Déscription de Denaea fournieri, sélacien nouveau du Marbre noir de Denée. Part 2 of Fournier, G. and Pruvost, P., Découverte d’un poisson nouveau dans le marbre noir de Denée. Bulletin de Academie Royal de Bruxelles Serie 5, 8, 213-218.

Robb III, A.J. 1992. Vertebrate fossils from the Lawrence Formation (Douglas Group, Virgilian, Upper Pennsylvanian) in Northeastern Kansas. Transactions of the Kansas Academy of Science 95(1/2), 129-138.View article

Roelofs, B., Barham, M., Mory, A.J. & Trinajstic, K. 2016. Late Devonian and Early Carboniferous chondrichthyans from the Fairfield Group, Canning Basin, Western Australia. Palaeontologia Electronica 19.1.4A,, 1-28.View article

Roelofs, B., Playton, T., Barham, M. & Trinajstic, K. 2015. Upper Devonian microvertebrates from the Canning Basin, Western Australia. Acta Geologica Polonica 65, 69-101.View article

Rosscoe, S.J. & Barrick, J.E. 2013. North American species of the conodont genus Idiognathodus from the Moscovian-Kasimovian boundary composite sequences and correlation of the Moscovian-Kasimovian boundary. New Mexico Museum of Natural History and Science, Bulletin 60, 354-371.

Soler-Gijón, R. 1997. Euselachian sharks from the Late Carboniferous of the Puertollano basin, Spain: biostratigraphic and palaeoenvironmental implications. Modern Geology 21, 137-169.

St. John, O.H. & Worthen, A.H. 1875, Descriptions of fossil fishes. Geological Survey of Illinois 6, 245-488.

Stahl, B.J. 1999. Chondrichthyes III. Holocephali, 1-164. In Schultze, H.-P. (ed.) Handbook of Palaeoichthyology 4. Verlag Dr. Friedrich Pfeil, München.

Wang, N-Z., Jin, F. & Wang, W. 2004. Early Carboniferous fishes (acanthodian, actinopterygians and Chondrichthyes) from the east sector of north Qilian Mountain, China - Carboniferous fish sequence from the east sector of north Qilian Mountain. Vertebrata Palasiatica 42, 89-110.

Williams, M.E. 1985. The ’cladodont level’ sharks of the Pennsylvanian black shales of central North America. Palaeontographica A 190(3/6), 83-158.

Zangerl, R. 1981. Chondrichthyes I. Paleozoic Elasmobranchii, 1-115. In Schultze, H.-P. (ed.) Handbook of Paleoichthyology 3A. Fischer, Stuttgart.

Zangerl, R. 1990. Two new stethacanthid sharks (Stethacanthidae, Symmoriida) from the Pennsylvanian of Indiana, U.S.A. Palaeontographica A 213, 115-141.

Zangerl, R. & Case, G.C. 1976. Cobelodus aculeatus (Cope), an anacanthous shark from Pennsylvanian black shales of North America. Palaeontographica A 154, 107-157.

Zidek, J. & Kietzke, K.E. 1993. Pre-Permian vertebrates of New Mexico, with remarks on some Early Permian specimens. New Mexico Museum of Natural History and Science, Bulletin 2, 1-10.