New vertebrate finds from the Khovanshchinian Regional Stage (lower uppermost Famennian, Upper Devonian) made an important contribution to our knowledge of the composition and distribution of vertebrate assemblages in Central Russia.
The remains of the latest arthrodires found for the first time in the East European Platform are described from the Voskresenskoye quarry (Lipetsk Region, Central Russia). Those occur within the deposits of the same basin, which had also been dwelled by tetrapods. The upper and uppermost Famennian non-groenlandaspidid arthrodires are known from only few localities in North America, Belgium and Poland. The separation of habitats of the tetrapod community and arthrodires in the Khovanshchinian basin may be due to local variations in the salinity level; these placoderms are possibly stenohaline marine dwellers. Environmental conditions of vertebrate communities in the Khovanshchinian Sea of the Moscow syneclise basin ranged from shallow-water brackish environment with still hydrodynamics and fast sedimentation to marine conditions with active hydrodynamics in the near-shore shallow waters. The osteolepiform genus Eusthenodon, recognized now from the Gorbachevo quarry (Tula Region, Central Russia), is characteristic of the Khovanshchinian vertebrate assemblage of Central Russia, but is also a marker genus of palaeotetrapod communities in Laurussia and East Gondwana.
Alekseev, A. S., Lebedev, O. A., Barskov, I. S., Barskova, M. I., Kononova, L. I. & Chizhova, V. A. 1994. On the stratigraphic position of the Famennian and Tournaisian fossil vertebrate beds in Andreyevka, Tula Region, Central Russia. Proceedings of the Geologists’ Association, 105, 41–52.
Alekseev, A. S., Kononova, L. I. & Nikishin, A. M. 1996. The Devonian and Carboniferous of the Moscow syneclise (Russian Platform): stratigraphy and sea-level changes. Tectonophysics, 268, 149–168.
https://doi.org/10.1016/S0040-1951(96)00229-6
Anderson, M. E., Hiller, N. & Gess, R. W. 1994. The first Bothriolepis-associated Devonian fish fauna from Africa. Suid-Afrikaanse Tydskrif vir Wetenskap, 90, 397–403.
Anderson, M. E., Long, J. A., Evans, F. J., Almond, J. E., Theron, J. N. & Bender, P. A. 1999. Biogeographic affinities of Middle and Late Devonian fishes of South Africa. Records of the Western Australia Museum, 57, 157–168.
Aristov, V. A. 1988. Devonskie konodonty Tsentralʹnogo devonskogo polya (Russkaya platforma) [Devonian Conodonts of the Central Devonian Field (Russian Platform)]. Nauka Publishers, Moscow, 119 pp. [in Russian].
Berg, L. S. 1940. Classification of fishes both recent and fossil. Trudy Zoologicheskogo Instituta, 5, 85–517 [in Russian].
Blieck, A., Clément, G., Blom, H., Lelièvre, H., Lukševičs, E., Streel, M., Thorez, J. & Young, G. C. 2007. The biostratigraphical and palaeogeographical framework of the earliest diversification of tetrapods (Late Devonian). In Devonian Events and Correlations (Becker, R. T. & Kirchgasser, W. T., eds), Geological Society, London, Special Publication, 278, 219–235.
https://doi.org/10.1144/SP278.10
Blom, H., Clack, J. A., Ahlberg, P. E. & Friedman, M. 2007. Devonian vertebrates from East Greenland: a review of faunal composition and distribution. Geodiversitas, 29, 119–141.
Branson, E. B. & Mehl, M. G. 1934. Conodonts from the Grassy Creek shale of Missouri. Missouri University Studies, 8, 171–259.
Breitbart, M., Hoare, A., Nitti, A., Siefert, J., Haynes, M., Dinsdale, E., Edwards, R., Souza, V., Rohwer, F. & Hollander, D. 2009. Metagenomic and stable isotopic analyses of modern freshwater microbialites in Cuatro Ciénegas, Mexico. Environmental Microbiology, 11, 16–34.
https://doi.org/10.1111/j.1462-2920.2008.01725.x
Burns, B. P., Goh, F., Allen, M. & Neilan, B. A. 2004. Microbial diversity of extant stromatolites in the hypersaline marine environment of Shark Bay, Australia. Environmental Microbiology, 6, 1096–1101.
Carr, R. K. & Jackson, G. L. 2009. Сhapter 5. The vertebrate fauna of the Cleveland Member (Famennian) of the Ohio Shale. In Guide to the Geology and Paleontology of the Cleveland Member of the Ohio Shale. 68th Annual Meeting of the Society of Vertebrate Paleontology, Cleveland, Ohio, October 15–18, 2008. Ohio Geological Survey Guidebook, 22, 1–17.
Clack, J. A. 2006. The emergence of early tetrapods. Paleogeography, Paleoclimatology, Paleoecology, 232, 167–189.
https://doi.org/10.1016/j.palaeo.2005.07.019
Cope, E. D. 1889. Synopsis of the families of Vertebrata. The American Naturalist, 23, 849–877.
https://doi.org/10.1086/275018
Denison, R. H. 1978. Placodermi. In Handbook of Paleoichthyology, Vol. 2. (Schultze, H. P., ed.). Gustav Fischer Verlag, Stuttgart, New York, 128 pp.
De Verneuil, E. 1845. Mollusques. In Geologie de la Russie d’Europe et des montagnes de l’Oural, 2 (Murchison, R. I., de Verneuil, E. & de Keyserling, A.), pp. 37–376. John Murray, Londres–Bertrand, Paris.
Druce, E. C. 1969. Devonian and Carboniferous conodonts from the Bonaparte Gulf Basin, Northern Australia and their use in international correlation. Bureau of Mineral Resources Bulletin, Geology and Geophysics, 69, 1–243.
Elliott, D. K., Johnson, H. G., Cloutier, R., Carr, R. K. & Daeschler, E. B. 2000. Middle and Late Devonian vertebrates of the western Old Red Sandstone Continent. In Palaeozoic Vertebrate Biochronology and Global Marine/Non-Marine Correlation (Blieck, A. & Turner, S., eds), Courier Forschungsinstitut Senckenberg, 223, 291–308.
Gatovsky, Yu. A. 2016. Conodont assemblages from the Upper Famennian–Lower Tournaisian boundary deposits in the central area of Russian Platform. In PALEOSTRAT-2016. Godichnoe sobranie (nauchnaya konferentsiya) sektsii paleontologii Moskovskogo obshchestva ispytatelej prirody i Moskovskogo otdeleniya Vserossijskogo Paleontologicheskogo obshchestva pri PIN RAS [Annual Meeting (Scientific Conference) of the Palaeontology Section of the Moscow Naturalists’ Society and the Moscow Branch of the Palaeontological Society of Russia affiliated to PIN RAS, Moscow, January 26–28, 2016. Abstracts Volume (Alekseev, A. S., ed.), pp. 24–26. Moscow [in Russian].
Gess, R. W. & Coates, M. I. 2015. High-latitude chondrichthyans from the Late Devonian (Famennian) Witpoort Formation of South Africa. Paläontologische Zeitschrift, 89, 147–169.
https://doi.org/10.1007/s12542-014-0221-9
Ginter, M. 2001. Chondrichthyan biofacies in the Late Permian of Utah and Nevada. Journal of Vertebrate Paleontology, 21, 714–729.
https://doi.org/10.1671/0272-4634(2001)021[0714:CBITLF]2.0.CO;2
Gurevich, K. Ya. 1972. Devonian and Early Carboniferous ostracods of the Volynʹ–Podolia margin of the Russian Platform and their stratigraphic importance. In Materialy po paleontolgii i stratigrafii nefte- i gazonosnykh rajonov zapadnoj chasti Ukrainskoj SSR [Materials on the Palaeontology and Stratigraphy of the Oil- and Gas-bearing Regions of the Western Part of the Ukrainian SSR] (Voloshina, A. M. & Pishvanova, L. S., eds), pp. 286–534. Nedra Publishers, Moscow [in Russian].
Hall, J. 1885. Lamellibranchiata 2. Dimyaria from the upper Helderberg, Hamilton, Portage, and Chemung groups. Natural History of New York, Palaeontology, 5, 269–561.
Janvier, P. & Clement, G. 2005. A new groenlandaspidid arthrodire (Vertebrata: Placodermi) from the Famennian of Belgium. Geologica Belgica, 8, 51–67.
Jarvik, E. 1952. On the fish-like tail in the ichthyostegid stegocephalians with descriptions of a new stegocephalian and a new crossopterygian from the Upper Devonian of East Greenland. Meddelelser om Grønland, 114, 1–90.
Johanson, Z. & Ritchie, A. 2000. Rhipidistians (Sarcopterygii) from the Hunter Siltstone (Late Famennian) near Grenfell, NSW, Australia. Fossil Record, 3, 111–136.
https://doi.org/10.5194/fr-3-111-2000
Kozmenko, A. S. 1911. Kratkoe predvaritel'noe soobshchenie o khode otsenochnykh gidrologicheskikh issledovanij v Tul′skoj gubernii za 1910 g. [Brief Preliminary Report on the Course of Estimative Hydrological Research in the Tula Governorate in 1910]. Tula Governorate Administration Printhouse, Tula, 133 pp. [in Russian].
Krupina, N. I. 1987. A new dipnoan from the Upper Devonian of the Tula Region. Paleontologicheskij Zhurnal, 3, 40–47 [in Russian].
Krupina, N. I. & Reisz, R. R. 1999. Reconstruction of dentition in hatchlings of Andreyevichthys epitomus, a Late Famennian dipnoan from Russia. Modern Geology, 24, 99–108.
Kulczycki, J. 1957. Upper Devonian fishes from the Holy Cross Mountains (Poland). Acta Palaeontologica Polonica, 2, 285–382.
Lebedev, O. A. 1984. The first find of a Devonian tetrapod in the USSR. Doklady Academii Nauk SSSR, 278, 1470–1473 [in Russian].
Lebedev, O. A. 1986. The new finds of the ichthyofauna and its value for the definition of the Devonian–Carboniferous boundary. In Geologiya i mineralʹnye resursy tsentralʹnykh rajonov Vostochno-Évropeyskoj platformy [The Geology and Mineral Resources of the Central Regions of the East-European Platform] (Zeisler, V. M., ed.), pp. 5–11. Nauka Publishers, Moscow [in Russian].
Lebedev, O. A. 1992. The latest Devonian, Khovanshchinian vertebrate community of Andreyevka-2 locality, Tula Region, Russia. In Fossil Fishes as Living Animals. Proceedings of II International Colloquium on the Study of Palaeozoic Fishes, Tallinn, 1989 (Mark-Kurik, E., ed.), Academia, 1, 265–272.
Lebedev, O. A. 2004. A new tetrapod Jakubsonia livnensis from the Early Famennian (Devonian) of Russia and palaeoecological remarks on the Late Devonian tetrapod habitats In The Second Gross Symposium “Advances of Palaeoichthyology” (Zelčs, V. & Lukševičs, E., eds), Acta Universitatis Latviensis, Earth and Environment Science, Special Publication, 679, 79–98.
Lebedev, O. A. 2013. Origin and evolution of tetrapod vertebrate communities (Devonian–Early Carboniferous). In Problemy evolyutsii biosfery [Problems of Biosphere Evolution] (Rozhnov, S. V., ed.), pp. 206–229. PIN RAS, Moscow [in Russian].
Lebedev, O. A., Lukševičs, E. & Zakharenko, G. V. 2010. Palaeozoogeographical connections of the Devonian vertebrate communities of the Baltica Province. Part II. Late Devonian. Palaeoworld, 19, 108–128.
https://doi.org/10.1016/j.palwor.2009.12.003
Lelièvre, H. 1982. Ardennosteus ubaghsi n. g., n. sp., Brachythoraci primitif (vertébré, placoderme) du Famennien d’Esneux (Belgique). Annales de la Société géologique de Belgique, 105, 1–7.
Linnaeus, C. 1758. Systema Naturae, 1. Leipzig, Engelman, 824 pp.
McCoy, F. 1848. On some new fossil fishes of the Carboniferous period. Annals and Magazine of Natural History, 2, 1–10, 115–133.
https://doi.org/10.1080/03745485809496133
Makhlaev, V. G. 1964. Usloviya osadkonakopleniya v Verkhnefamenskom bassejne Russkoj platformy [Sedimentary Conditions in the Upper Famennian Basin of the Russian Platform]. Nauka, Moscow, 227 pp. [in Russian].
Menning, M., Alekseev, A. S., Chuvashov, B. I., Davydov, V. I., Devuyst, F. X., Forke, H. C. & Jin, Y. G. 2006. Global time scale and regional stratigraphic reference scales of central and west Europe, east Europe, Tethys, south China, and North America as used in the Devonian–Carboniferous–Permian Correlation Chart 2003 (DCP 2003). Palaeogeography, Palaeoclimatology, Palaeoecology, 240, 318–372.
https://doi.org/10.1016/j.palaeo.2006.03.058
Müller, J. 1845. Über den Bau und die Grenzen der Ganoiden und über das natürliche System der Fische. Archiv für Naturgeschichte, 11, 91–141.
Nalivkin, B. V. 1947. Class Lamellibranchiata. Lamellibranchiates. In Atlas rukovodyashchikh form iskopayemykh faun SSSR, tom 3, Devonskaya sistema [Atlas of the Index Fossils of Extinct Faunas of the USSR, Vol. 3, Devonian System] (Nalivkin, D. V., ed.), pp. 134–147. Moscow–Leningrad [in Russian].
Nalivkin, B. V. 1953. Stratigraphic importance of pelecypods for the Devonian deposits of the Russian Platform. In Devon Russkoj Platformy [Devonian of the Russian Platform] (Ivanchuk, P. K., ed.), pp. 248–259. Gostoptekhizdat Publishers, Leningrad–Moscow [in Russian].
Naumova, S. N. 1953. Spore assemblages in the Upper Devonian of the Russian Platform and their importance for stratigraphy. Trudy Instituta Geologicheskikh Nauk AN SSSR, Geologiya, 143, 3–202 [in Russian].
Newberry, J. S. 1873. Paleontology. Descriptions of fossil fishes. Geological Survey of Ohio, 1, 245–255.
Obrucheva, O. P. 1956. Dinichthyid remains from the Upper Devonian of USSR. Doklady Akademii Nauk SSSR, 108, 333–336 [in Russian].
Reutlinger, E. A. 1960. Characterisation of the Ozerki and Khovanshchinian Beds by microscopic organic remains (central part of the Russian Platform). Trudy Geologicheskogo Instituta AN SSSR, 14, 136–172.
Rhodes, F. H. T., Austin, R. L. & Druce, E. C. 1969. British Avonian (Carboniferous) conodont faunas, and their value in local and intercontinental correlation. Bulletin of the British Museum (Natural History), Supplements, 5, 1–313.
Rodionova, G. D., Umnova, V. T., Kononova, L. I., Ovnatanova, N. S., Rzhonsnitskaya, M. A. & Fedorova, T. I. 1995. Devon Voronezhskoj anteklizy i Moskovskoj sineklizy [The Devonian of the Voronezh Anteclise and Moscow Syneclise]. Rosgeolfond, Moscow, 265 pp. [in Russian].
Romer, A. S. 1955. Herpetichthyes, Amphibioidei, Choanichthyes or Sarcopterygii? Nature, 176, 126.
https://doi.org/10.1038/176126a0
https://doi.org/10.1038/176126b0
Rzhonsnitskaya, M. A. & Kulikova, V. F. (eds). 1990. Reshenie mezhvedomstvennogo regionalʹnogo stratigraficheskogo soveshchaniya po srednemu i verkhnemu paleozoyu Russkoj platformy s regionalʹnymi stratigraficheskimi skhemami. Devonskaya sistema [Decisions of the Interdepartmental Regional Stratigraphic Session (IRSS) on the Upper and Middle Palaeozoic of the Russian Platform with Regional Stratigraphic Charts (Leningrad, 1988). Devonian System]. VNIGRI, Leningrad, 60 pp. [in Russian].
Samoilova, R. B. 1954. On the characteristics of the Upper Devonian and Lower Carboniferous by microfauna. Byulletenʹ Moskovskogo Obshchestva Ispytatelej Prirody, Geologiya, 29, 92 [in Russian].
Samoilova, R. B. 1979. Key ostracod assemblages of the Lower Carboniferous of the Moscow Syneclise. In Stratigrafiya, paleontologiya i paleogeografiya karbona Moskovskoj sineklizy [Stratigraphy, Palaeontology and Palaeogeography of the Carboniferous of the Moscow Syneclise] (Makhlina, M. Kh. & Schick, S. M., eds), pp. 76–87. Geolfond RSFSR, Moscow [in Russian].
Shulga, P. L. 1962. Materials of the Permanent Stratigraphic Commission of the Interdepartmental Stratigraphic Commitee on the Devonian of USSR. Byulletenʹ Mezhvedomstvennogo Stratigraficheskogo Komiteta, 4, 70–71 [in Russian].
Stensiö, E. A. 1944. Contributions to the knowledge of the vertebrate fauna of the Silurian and Devonian of Western Podolia. Part II. Notes on two arthrodires from the Downtonian of Podolia. Arkiv för Zoologi, 35, 1–83.
Stensiö, E. A. 1963. Anatomical studies on the arthrodiran head. Part 1. Preface, geological and geographical distribution, the organization of the head in the Dolichothoraci, Coccosteomorphi and Pachyosteomorphi. Taxonomic appendix. Kungliga Svenska Vetenskapsakademiens Handlingar, 9, 1–419.
Thomas, L. A. 1949. Devonian–Mississippian formations of southeast Iowa. Geological Society of America Bulletin, 60, 403–438.
https://doi.org/10.1130/0016-7606(1949)60[403:DFOSI]2.0.CO;2
Vanuxem, L. 1842. Geology of New York, Part III. Comprising the Survey of the Third Geologic District, Albany, NY. Natural History of New York Division, Geology, 3, 3–307.
Venyukov, P. N. 1886. Fauna of the Devonian system of the north-west and Central Russia. Trudy Sankt-Peterburgskogo Obshchestva Estestvoispytatelej, 17, I–XVI + 417–708 [in Russian].
Vézina, D. 1990. Les Plourdosteidae fam. nov. (Placodermi, Arthrodira) et leurs relations phylétiques au sein des Brachythoraci. Canadian Journal of Earth Sciences, 27, 677–683.
https://doi.org/10.1139/e90-065
Woodward, A. S. 1891. Catalogue of the Fossil Fishes in the British Museum (Natural History). Pt. II. Elasmobranchii (Acanthodii), Holocephali, Ichthyodorulites, Ostracodermi, Dipnoi, and Teleostomi (Crossopterygii and Chondrostean Actinopterygii). Taylor & Francis, London, xliv+567 pp.
Young, G. C. & Turner, S. 2000. Devonian microvertebrates and marine-nonmarine correlation in East Gondwana: overview. In Palaeozoic Vertebrate Biochronology and Global Marine/Non-Marine Correlation (Blieck, A. & Turner, S., eds), Courier Forschungsinstitut Senckenberg, 223, 453–470.
Zhu, M., Ahlberg, P. E., Zhao, W. & Jia, L. 2002. First Devonian tetrapod from Asia. Nature, 420, 760–761.
https://doi.org/10.1038/420760a
Ziegler, W. & Sandberg, C. A. 1990. The late Devonian Standard Conodont Zonation. Courier Forschungsinstitut Senckenberg, 121, 3–115.
