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Squamation of the Permian actinopterygian Toyemia Minich, 1990: evenkiid (Scanilepiformes) affinities and implications for the origin of polypteroid scales
Published in: Bulletin of Geosciences, volume 97, issue 2; pages: 235 - 259; Received 11 August 2021; Accepted in revised form 28 February 2022; Online 5 June 2022
Keywords: ganoid scales, lepidotrichia, palaeohistology, Osteichthyes, Scanilepiformes, Permian, European Russia,
Abstract
Toyemia Minich, 1990, is a widespread member of late Permian freshwater fossil assemblages of European Russia. The scales and lepidotrichia hemisegments of this taxon show general resemblance to the palaeoniscoid-type ‘ganoid’ scales of most Palaeozoic actinopterygians, but differ from these in some morphological features. Architecture of the vascular system is more complex: part of the canals (the wider ones) are located within the bone layer, constituting a complicated three-dimensional pattern, while the thinner canals usually lie on the border of the bone and the dentine layers (with the canal lumen usually surrounded by dentine but positioned in the basalmost part of the odontode), forming a regular network, especially in the posterior and ventral part of a scale. Whereas in most basal actinopterygians, vascular canals only occur underneath the sculptural ridges of the free field, scales of Toyemia possess wide, sinuous canals underneath the depressed field, opening to its surface in large pores. These and further features of the squamation (vascular canals cased in concentric layers of bone reminiscent of osteons and surrounded by numerous osteocyte lacunae; the presence of true secondary osteons; the shape of articulating elements and the general scale outline; the ornamentation of the free field, mainly consisting of concentric rolls; etc.) are similar to representatives of the Triassic order Scanilepiformes, especially to the family Evenkiidae, suggesting that Toyemia should be placed within this family. An enigmatic ’circular structure’ of interwoven collagen fibers, first observed in Toyemia scales, is reminiscent of polypterid elasmodine. In several aspects, scales of Toyemia appear to represent the ancestral condition of the polypteroid scale.References
Aldinger, H. 1937. Permische Ganoidfische aus Ostgrönland. Meddelelser om Gronland, 102(3), 1-392.
Arefiev, M.P., Golubev, V.K., Balabanov, Yu.P., Karasev, E.V., Minikh, A.V., Minikh, M.G., Molostovskaya, I.I., Yaroshenko, O.P.& Zhokina-Naumcheva, M.A. 2015. Type and reference sections of the Permian-Triassic continental sequences of the East European Platform: main isotope, magnetic, and biotic events. 104 pp. PIN RAS, Moscow.
Bakaev, A.S. & Bulanov, V.V. 2021. The squamation morphology and stratigraphic distribution of Isadia arefievi A. Minich (Actinopterygii, Eurynotoidiformes). Paleontological Journal 55(2), 205-216.
Bakaev, A. & Kogan I. 2020. A new species of Burguklia (Pisces, Actinopterygii) from the Middle Permian of the Volga Region (European Russia). PalZ 94, 93-106.
Bakaev, A.S., Kogan, I., Silantiev, V.V., Golubev, V.K. & Schneider, J.W. 2017. Changes in Middle and Late Permian ichthyofaunas of European Russia - causes, significance, global correlation. 88. Jahrestagung der Paläontologischen Gesellschaft, Münster, 26.-30. März 2017, Münstersche Forschungen zur Geologie und Paläontologie, 109, 27.
Bakaev, A.S., Kogan, I. & Yankevich, D.I. 2020. On the validity of names of some Permian actinopterygians from European Russia. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 296(3), 305-316.
Berg, L.S. 1941. Lower Triassic fishes of the Tunguska Coal basin, Yenisei, Siberia. Bulletin of the USSR Academy of Sciences. Biological series 1941, 458-474.
Burrow, C.J. 1994. Form and function in scales of Ligulalepis toombsi Schultze, a palaeoniscoid from the Early Devonian of Australia. Records of the South Australian Museum 27, 175-185.
Chen, D., Janvier, P., Ahlberg, P.E. & Blom, H. 2012. Scale morphology and squamation of the Late Silurian osteichthyan Andreolepis from Gotland, Sweden. Historical Biology: An International Journal of Paleobiology 24(4), 411-423.
Choo, B. 2011. Revision of the actinopterygian genus Mimipiscis (= Mimia) from the Upper Devonian Gogo Formation of Western Australia and the interrelationships of the early Actinopterygii. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 11, 1-28.
Choo, B., Long, J.A. & Trinajstic, K. 2009. A new genus and species of basal actinopterygian fish from the Upper Devonian Gogo Formation of Western Australia. Acta Zoologica (Stockholm) 90, 194-210.
Choo, B., Zhu, M., Qu, Q., Yu, X., Jia, L. & Zhao, W. 2017. A new osteichthyan from the Late Silurian of Yunnan, China. PloS One 12(3), e0170929.
Cope, E.D. 1887. General notes: geology and palaeontology. IV. American Naturalist 21(11), 1016-1019.
Cui, X., Qiao, T. & Zhu, M. 2019. Scale morphology and squamation pattern of Guiyu oneiros provide new insights into early osteichthyan body plan. Scientific Reports 9, 4411.
Cuvier, G. 1829. Le regne animal distribué d’apres son organisation, pour servir de base a l’histoire naturelle des animaux, et d’introduction a l’anatomie comparée. Nouvelle Édition, Tome II. 406 pp. Déterville & Crochard, Paris.
Eichwald, E. 1860. Lethaea Rossica ou Paléontologie de la Russie, Vol. 1. 1657 pp. Schweizerbart, Stuttgart.
Esin, D.N. 1989. Scales of polypterids (?) (Actinopterygii) in Upper Permian deposits of the East European Platform, and the problem of the geological history of this group [Cheshui mnogopiorov (?) (luchepiorye) v verkhnepermskikh otlozheniyakh Vostochno-Evropeyskoy platformy i problema geologicheskoy istorii etoy gruppy]. Bulletin of Moscow Society of Naturalists, Geological section 64(5), 128-129. [in Russian]
Esin, D.N. 1990. The scale cover of Amblypterina costata (Eichwald) and the paleoniscid taxonomy based on isolated scales. Paleontological Journal 1990(2), 90-98.
Esin, D.N. 1995. Late Permian palaeoniscids of the European part of Russia [Posdnepermskiye paleoniscidy Evropeyskoj casti Rossii]. 250 pp. Ph.D. thesis, Lomonossov State University, Moscow, Russia. [in Russian]
Esin, D.N. & Mashin, V.L. 1996. Ichthyoliths [Ikhtiolity], 270-293. In Esaulova, N.K. & Lozovsky, V.R. (eds) Stratotypes and reference sections of the Upper Permian of Povolzhye and Prikamye [Stratotipy i opornye razrezy verkhney permi Povolzh’ya i Prikam’ya]. Ecocenter, Kazan. [in Russian]
Francillon-Viellot, H., Buffrénil, V. de, Castanet, J., Géraudie, J., Meunier, F.J., Sire, J.Y., Zylberberg, L. & Ricqles, A. de 1990. Microstructure and mineralization of vertebrate skeletal tissues, 471-530. In Carter, J.G. (ed.) Skeletal biomineralization: patterns, processes and evolutionary trends. Van Nostrand Reinhold, New York.
Franeck, F. 2013. Micropalaeontology of isolated vertebrate remains from the Trassic of Kyrgyzstan (Madygen-Fm.). 61 pp. Bachelor thesis, TU Bergakademie Freiberg, Germany.
Gardiner, B.G. 1967. Further notes on palaeoniscoid fishes with a classification of the Chondrostei. Bulletin of the British Museum (Natural History), Geology 14(5), 143-206.
Gayet, M. & Meunier, F.J. 1992. Polyptériformes (Pisces, Cladistia) du Maastrichtien et du Paléocene de Bolivie. Geobios, Mém. Spéc. 14, 159-168.
Gayet, M., Meunier F.J. & Werner, C. 2002. Diversification in Polypteriformes and special comparison with the Lepisosteiformes. Palaeontology 45, 361-376.
Geerlink, P.J. & Videler, J.J. 1987. The relation between structure and bending properties of teleost fin rays. Netherlands Journal of Zoology 37(1), 59-80.
Gemballa, S. & Bartsch, P. 2002. Architecture of the integument in lower teleostomes: functional morphology and evolutionary implications. Journal of Morphology 253, 290-309.
Giles, S., Coates, M.I., Garwood, R.J., Brazeau, M.D., Atwood, R. & Johanson, Z., Friedman, M. & Ruta, M. 2015. Endoskeletal structure in Cheirolepis (Osteichthyes, Actinopterygii), an early ray-finned fish. Palaeontology 58, 849-870.
Giles, S., Xu, G.-H., Near, T.J. & Friedman, M. 2017. Early members of ’living fossil’ lineage imply later origin of modern ray-finned fishes. Nature 549, 265-268.
Goodrich, E.S. 1907. On the scales of fish, living and extinct, and their importance in classification. Proceedings of the Zoological Society of London 1907, 751-774.
Goodrich, E.S. 1930. Studies on the structure and development of Vertebrates. 837 pp. Macmillan, London.
Grandstaff, B.S., Smith, J.B., Lamanna, M.C., Lacovara, K.J. & Abdel-Ghani, M.S. 2012. Bawitius, gen. nov., a giant polypterid (Osteichthyes, Actinopterygii) from the Upper Cretaceous Bahariya Formation of Egypt. Journal of Vertebrate Paleontology 32(1), 17-26.
Gross, W. 1956. Über Crossopterygier und Dipnoer aus dem baltischen Oberdevon im Zusammenhang einer vergleichenden Untersuchung des Porenkanalsystems paläozoischer Agnathen und Fische. Kungliga Svenska Vetenskapsakademiens Handlingar, 4. Ser. 5(6), 1-140.
Gross, W. 1968. Fragliche Actinopterygier-Schuppen aus dem Silur Gotlands. Lethaia 1, 184-218.
Ivanov, A.O. 2005. Early Permian chondrichthyans of the Middle and South Urals. Revista Brasileira de Paleontologia 8(2), 127-138.
Ivanov, A.O., Bakaev, A.S., Nestell, M.K. & Nestell, G.P. 2021. Fish microremains from the Cutoff Formation (Roadian, Middle Permian) of the Guadalupe Mountains, West Texas, USA. Micropaleontology 67, 365-402.
Jerve, A., Qu, Q., Sanchez, S., Blom, H. & Ahlberg, P.E. 2016. Three-dimensional paleohistology of the scale and median fin spine of Lophosteus superbus (Pander 1856). PeerJ e2521, 1-29.
Kerr, T. 1952. The scales of primitive living actinopterygians. Proceedings of the Zoological Society of London 122, 55-78.
Kogan, I. & Bakaev, A.S. 2019. Permian and Triassic scanilepiforms (Actinopterygii) of Eastern Europe and Northern Asia and their polypterid affinity. 90th annual meeting of the Paläontologische Gesellschaft, 15.-18.9.2019, Munich, Germany, Abstracts, 84.
Krotow, B.P. 1904. Fishes of the Permian deposits of Russia [Ryby permskikh otlozheniy Rossii]. Proceedings of the Society of Naturalists at the Imperial Kazan University translation 38(3), 1-40. [in Russian]
Lehman, J.P. 1979. Le genre Scanilepis Aldinger du Rhetien de la Scanie (Suede). Bulletin of the Geological Institution of the University of Uppsala 8, 113-125.
Liu, G.-B. & Shen, C.-M. 2006. A new progressive palaeoniscoid, Mizhilepis from the Upper Triassic in Mizhi of North Shaanxi. Acta Palaeontologica Sinica 45, 514-522.
Long, J.A., Choo, B. & Young, G.C. 2008. A new basal actinopterygian from the Middle Devonian Aztec Siltstone of Antarctica. Antarctic Science 20(4), 393-412.
Lu, J., Giles, S., Friedman, M., den Blaauwen, J.L. & Zhu, M. 2016. The oldest actinopterygian highlights the cryptic early history of the hyperdiverse ray-finned fishes. Current Biology 26(12), 1602-1608.
Märss, T. 2006. Exoskeletal ultrastructure of early vertebrates. Journal of Vertebrate Paleontology 26(2), 235-252.
Minikh, A.V. 1995. New actinopterygian fishes of the Tatarian stage from the Sukhona River basin and Southern Cis-Urals [Novye luchepiorye ryby iz tatarskogo yarusa basseina r. Suchony i Yuzhnogo Priural’ya]. 18 pp. Saratov Unviersity Press. Saratov. Deposited in VINITI on 10.05.1995 under No. 1306 - V95. [in Russian]
Minikh, A.V. & Minikh, M.G. 2009. Ichthyofauna of the Permian of European Russia [Ikhtiofauna permi Evropeyskoy Rossii]. 243 pp. Nauka, Saratov. [in Russian]
Minikh, M.G. & Minikh, A.V. 1990. Revision of some palaeoniscids and new fishes from the Upper Permian of the East European Platform, and possibilities of using them stratigraphically [Revisiya nekotorykh paleoniskov i novye ryby iz verkhney permi Vostochno-Evropeyskoy platformy i vozmozhnosti ikh ispolzovaniya v stratigrafii], 84-104. In Aleshechkin, O.I. (ed.) Problems of the geology of South Urals and Lower Volga Region [Voprosy geologii Yuzhnogo Urala i Nizhnego Povolzh’ya]. SarGU Press, Saratov. [in Russian]
Mondéjar-Fernández, J. 2018. On cosmine: its origins, biology and implications for sarcopterygian interrelationships. Cybium, 42(1), 41-65.
Mondéjar-Fernández, J. & Clément, G. 2012. Squamation and scale microstructure evolution in the Porolepiformes (Sarcopterygii, Dipnomorpha) based on Heimenia ensis from the Devonian of Spitsbergen. Journal of Vertebrate Paleontology 32(2), 267-284.
Mondéjar-Fernández, J. & Meunier, F.J. 2021. New histological information on Holoptychius Agassiz, 1839 (Sarcopterygii, Porolepiformes) provides insights into the palaeoecological implications and evolution of the basal plate of the scales of osteichthyans. Historical Biology 33(10), 2276-2288.
Orvig, T. 1978. Microstructure and growth of the dermal skeleton in fossil actinopterygian fishes: Birgeria and Scanilepis. Zoologica Scripta 7, 33-56.
Pander, C.H. 1856. Monographie der fossilen Fische des silurischen Systems der Russisch-Baltischen Gouvernements. 91 pp. Kaiserliche Akademie der Wissenschaften, St. Petersburg.
Qu, Q., Sanchez, S., Blom, H., Tafforeau, P. & Ahlberg, P.E. 2013a. Scales and tooth whorls of ancient fishes challenge distinction between external and oral “teeth”. PLoS One 8(8), e71890.
Qu, Q., Zhu, M. & Wang, W. 2013b. Scales and dermal skeletal histology of an early bony fish Psarolepis romeri and their bearing on the evolution of rhombic scales and hard tissues. PLoS One 8(4), e61485.
Qu, Q., Haitina, T., Zhu, M. & Ahlberg, P.E. 2015. New genomic and fossil data illuminate the origin of enamel. Nature 526, 108-111.
Qu, Q., Sanchez, S., Zhu, M., Blom, H. & Ahlberg, P.E. 2017. The origin of novel features by changes in developmental mechanisms: ontogeny and three-dimensional microanatomy of polyodontode scales of two early osteichthyans. Biological Reviews 92(2), 1189-1212.
Richter, M. & Smith, M.M. 1995. A microstructural study of the ganoine tissue of selected lower vertebrates. Zoological Journal of the Linnean Society 114, 173-212.
Romano, C., Koot, M.B., Kogan, I., Brayard, A., Minikh, A.V., Brinkmann, W., Bucher, H. & Kriwet, J. 2016. Permian-Triassic Osteichthyes (bony fishes): Diversity dynamics and body size evolution. Biological Reviews 91(1), 106-147.
Schaeffer, B. 1967. Late Triassic fishes from the Western United States. Bulletin of the American Museum of Natural History 135, 285-342.
Schaeffer, B. & McDonald, N.G. 1978. Redfieldiid fishes from the Triassic-Liassic Newark Supergroup of eastern North America. Bulletin of the American Museum of Natural History 159, 131-173.
Schultze, H.-P. 1966. Morphologische und histologische Untersuchungen an Schuppen mesozoischer Actinopterygier (Übergang von Ganoid- zu Rundschuppen). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 126(3), 232-314.
Schultze, H.-P. 1968. Palaeoniscoidea-Schuppen aus dem Unterdevon Australiens und Kanadas und aus dem Mitteldevon Spitzbergens. Bulletin of the British Museum (Natural History) Geology 16, 342-368.
Schultze, H.-P. 2016. Scales, enamel, cosmine, ganoine, and early osteichthyans. Comptes Rendus Palevol 15(1-2), 83-102.
Schultze, H.-P. 2018. Hard tissues in fish evolution: history and current issues. Cybium 42(1), 29-39.
Selezneva, A.A. 1985. Evenkia - Ancestor of Polypterus (Actinopterygii). Paleontological Journal 19, 1-6.
Sire, J.-Y., Donoghue, P.C.J. & Vickaryous, M.K. 2009. Origin and evolution of the integumentary skeleton in non-tetrapod vertebrates. Journal of Anatomy 214, 409-440.
Su, D.-Z. 1978. A new Triassic palaeoniscoid fish from Fukang, Xinjiang. Memoirs of Institute of Vertebrate Palaeontology and Palaeoanthropology, Academia Sinica 13, 55-59. [in Chinese]
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. Pfeil, München.
Trinajstic, K. 1999a. Scale morphology of the Late Devonian palaeoniscoid Moythomasia durgaringa Gardiner and Bartram, 1977. Alcheringa 23, 9-19.
Trinajstic, K. 1999b. Palaeoniscoid scales from the Late Devonian of Western Australia. Records of the Western Australian Museum Supplement 57, 93-106.
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, 27-57.
Witzmann, F. 2011. Morphological and histological changes of dermal scales during the fish-to-tetrapod transition. Acta Zoologica 92(3), 281-302.
Xu, G.-H. & Gao, K.-Q. 2011. A new scanilepiform from the Lower Triassic of northern Gansu Province, China, and phylogenetic relationships of non-teleostean Actinopterygii. Zoological Journal of the Linnean Society 161, 595-612.
Yakovlev, V.N. 1973. Freshwater actinopterygians of the Mesozoic of Asia [Presnovodnye luchepiorye mezozoya Asii]. 28 pp. Author’s summary of Ph.D. thesis, Moscow State University, Moscow, USSR. [in Russian]
Zhu, M., Yu, X., Wang, W., Zhao, W. & Jia, L. 2006. A primitive fish provides key characters bearing on deep osteichthyan phylogeny. Nature 441, 77-80.
Zhu, M., Wang, W. & Yu, X. 2010. Meemannia eos, a basal sarcopterygian fish from the Lower Devonian of China-expanded description and significance, 199-214. In Elliot, D.K., Maisey, J.G., Yu, X. & Miao, D. (eds), Morphology, phylogeny and paleobiogeography of fossil fishes. Pfeil, München.
Zylberberg, L. & Meunier, F.J. 2013. A revisited histological study of the finrays of Polypterus senegalus (Cladistia, Polypteridae). Comptes Rendus Palevol 12, 203-210.
Zylberberg, L., Meunier, F.J. & Laurin, M. 2010. A microanatomical and histological study of the postcranial dermal skeleton in the Devonian sarcopterygian Eusthenopteron foordi. Acta Palaeontologica Polonica 55(3), 459-470.
Zylberberg, L., Meunier, F.J. & Laurin, M. 2016. A microanatomical and histological study of the postcranial dermal skeleton of the Devonian actinopterygian Cheirolepis canadensis. Acta Palaeontologica Polonica 61(2), 363-376.