The Devonian psammosteid agnathan Psammosteus livonicus Obruchev is revised using analysis of fossil material from collections in Russia, Latvia, Lithuania and Estonia that allowed the diagnosis and description of this species to be substantially extended. Psammosteus livonicus is similar to other species of Psammosteus in the characters of the dorsal plate, cyclomorial tesserae and stenobasal shape of the branchial plates. The study of the morphology of the branchial plates and tesserae (genus-level features) with consideration of species-level features of the ornamentation proves that the establishment of the genus Oredezhosteus Moloshnikov is baseless. The species Psammolepis aerata Obruchev in Halstead Tarlo 1965 and Oredezhosteus kuleshovi Moloshnikov 2009 are included as junior synonyms into the revised species based on the ornamentation features.
Agassiz, L. 1840. Tableau général des Poissons fossiles. In Recherches sur les Poissons fossiles, Neuchâtel, Soleure, 1, 32–49.
Agassiz, L. 1844. Monographie des Poissons fossiles du vieux grés rouge ou système Dévonien (Old Red Sandstone) des Isles Britanniques et de Russie. Neuchâtel, 188 pp.
Berg, L. S. 1940. Sistema ryboobraznykh i ryb, nyne zhivushchikh i iskopaemykh [Classification of fishes, both recent and fossil]. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR [Proceedings of the Zoological Institute of the Academy of Sciences of the USSR], 5, 1–517 [in Russian, partly in English, pp. 346–517].
Berg, L. S. 1955. Sistema ryboobraznykh i ryb, nyne zhivushchikh i iskopaemykh [Classification of fishes, both recent and fossil]. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR [Proceedings of the Zoological Institute of the Academy of Sciences of the USSR], 20, 1–286 [in Russian].
Brotzen, F. 1936. Beiträge zur Vertebratenfauna des westpodolischen Silurs und Devons. I. Protaspis arnelli n. sp. und Brachipteraspis n. gen. latissima Zych. Svenska Vetenskapsakad. Arkiv för Zoologi, 28A, 1–52.
Elliott, D. & Mark-Kurik, E. 2005. A review of the lateral line sensory system in psammosteid heterostracans. Revista Brasileira de Paleontologia, 8, 99–108.
http://dx.doi.org/10.4072/rbp.2005.2.02
Elliott, D. K., Mark-Kurik, E. & Daeschler, E. B. 2004. A revision of Obruchevia (Psammosteida: Heterostraci) and a description of a new obrucheviid from the Late Devonian of the Canadian Arctic. Acta Universitatis Latviensis, 679, 22–45.
Esin, D., Ginter, M., Ivanov, A., Lebedev, O., Lukševičs, E., Avkhimovich, V., Golubtsov, V. & Petukhova, L. 2000. Vertebrate correlation of the Upper Devonian and Lower Carboniferous on the East European Platform. Courier Forschungsinstitut Senckenberg, 223, 341–359.
Glinskiy, V. 2013. Topograficheskaya izmenchivost¢ ornamenta devonskikh beschelyustnykh – psammosteid [Topographic variability of the ornamentation of Devonian jawless vertebrates – psammosteids]. In Sovremennaya paleontologiya: Klassicheskie i novejshie metody. Desyataya Vserossijskaya nauchnaya shkola molodykh uchenykh-paleontologov [Modern Paleontology: Classical and Newest Methods. The Tenth All-Russian Scientific School for Young Scientists in Paleontology. Abstracts] (Lopatin, A. V., Parkhaev, P. Yu. & Rozanov, A. Yu., eds), pp. 15–16. PIN RAN Press, Moscow [in Russian].
Glinskiy, V. & Ivanov, A. 2015. The assemblages of psammosteid agnathans from the Middle–Late Devonian of the Andoma Hill (Russia). In IGCP596–SDS Symposium (Brussels, September 2015): Climate Change and Biodiversity Patterns in the Mid-Palaeozoic, Abstracts (Mottequin, B., Denayer, J., Königshof, P., Prestianni, C. & Olive, S., eds), Strata, série 1: Communications, 16, 57–59.
Gross, W. 1935. Histologische Studien am Aussenskelett fossiler Agnathen und Fische. Palaeontographica, 83A, 1–60.
Gross, W. 1937. Die Wirbeltiere des rheinischen Devons. Abhandlungen der Preußischen Geologischen Landesanstalt, 176, 1–83.
Gross, W. 1942. Die Fischfaunen des baltischen Devons und ihre biostratigraphische Bedeutung. Korrespondenblatt des Naturforscher–Vereins zu Riga, 64, 373–436.
Gross, W. 1963. Drepanaspis gemuendenensis Schlüter Neuuntersuchung. Palaeontographica, 121, 1–15.
Halstead, L. B. 1969. Calcified tissues in the earliest vertebrates. Calcified Tissue Research, 3, 107–124.
Halstead, L. B. 1993. Agnatha. In The Fossil Record 2 (Benton, M. J., ed.), рр. 573–581. Chapmann & Hall, London.
Halstead Tarlo, L. B. 1964. Psammosteiformes (Agnatha) – A review with descriptions of new material from the Lower Devonian of Poland. I. General part. Palaeontologia Polonica, 13, 1–135.
Halstead Tarlo, L. B. 1965. Psammosteiformes (Agnatha) – A review with descriptions of new material from the Lower Devonian of Poland. II. Systematic part. Palaeontologia Polonica, 15, 1–167.
Hekker, R. F. & Philippova, M. F. 1935. Razrez po ruch¢yu Tesovoj, pritoku Oredezha [Geological section on the Creek Tesovoj, River Oredezh affluent]. Trudy Leningradskogo geologo-gidrogeodezicheskogo tresta, 5–7, 23–36 [in Russian].
Ivanov, A. & Lebedev, O. 2011. Devonian Vertebrate Localities in the Luga River Basin (Leningrad Region, Russia). Guidebook for the Field Trip. St. Petersburg University Press, St. Petersburg, 37 pp.
Ivanov, A., Lukševičs, E., Stinkulis, G., Tovmasjan, K., Zupiņš, I. & Zabele, A. 2005. Devonian stratigraphy and vertebrate fauna of the Andoma Hill section (Onega Lake, Russia). In Middle Paleozoic Vertebrates of Laurussia: Relationships with Siberia, Kazakhstan, Asia and Gondwana. Ichthyolith Issues, Special Publication, 9, 17–21.
Janvier, P. 1996. Early Vertebrates. Clarendon Press, Oxford, 393 pp.
Johanson, Z., Smith, M., Kearsley, A., Pilecki, P., Mark-Kurik, E. & Howard, C. 2013 Origins of bone repair in the armour of fossil fish: response to a deep wound by cells depositing dentine instead of dermal bone. Biology Letters, 9, 20130144.
Karatajūtė-Talimaa, V. N. 1966. Botriolepidy shventoiskogo gorizonta Pribaltiki [Bothriolepids of the Šventoji Horizon of the East Baltic area]. In Paleontologiya i stratigrafiya Pribaltiki i Belorussii [Palaeontology and Stratigraphy of the Baltic and Byelorussia], Vol. 1(6) (Grigelis, A. A., ed.), pp. 191–279. Mintis, Vilnius [in Russian, with English summary].
Keating, J. N., Marquart, C. L. & Donoghue, P. C. J. 2015. Histology of the heterostracan dermal skeleton: insight into the origin of the vertebrate mineralised skeleton. Journal of Morphology, 276, 657–680.
Kiaer, J. 1932. The Downtonian and Devonian vertebrates of Spitsbergen. IV. Suborder Cyathaspida. A preliminary report edited by A. Heintz. Skrifter Svalbard Ishavet, 52, 1–26.
Kuršs, V. M., Viiding, H. A. & Mark-Kurik, E. Yu. 1981. Amata Regional Stage. In Devon i karbon Pribaltiki [Devonian and Carboniferous of the Baltic Area] (Sorokin, V. S., ed.), pp. 157–167. Zinātne, Rīga [in Russian].
Lankester, E. R. 1868. On the discovery of the remains of cephalaspidian fishes in Devonshire and Cornwall; and on the identity of Steganodictyum, M’Coy, with genera of those fishes. Quarterly Journal of the Geological Society of London, 24, 546–547.
Lukševičs, E. V., Ivanov, A. O. & Zupiņš, I. A. 2012. Kompleksy devonskikh pozvonochnykh Andomskoj gory i korrelyatsiya s razrezami Glavnogo devonskogo polya [Assemblages of Devonian vertebrates from Andoma Hill and correlation with sections of the Main Devonian Field]. In Paleozoj Rossii: regional¢naya stratigrafiya, paleontologiya, geo- i biosobytiya. Materialy III Vserossijskogo soveshchaniya [Paleozoic of Russia: Regional Stratigraphy, Palaeontology, Geo- and Bioevents. Materials of the Third All-Russian Conference] (Zhamojda, A. I., ed.), pp. 128–130. VSEGEI, St. Petersburg [in Russian].
Lyarskaya, L. 1972. A classification of Devonian vertebrate localities of Latvia. Eesti NSV Teaduste Akadeemia Toimetised, Keemia, Geoloogia, 21, 259–268.
Mark-Kurik, E. 1968. New finds of the psammosteids (Heterostraci) in the Devonian of Estonia and Latvia. Eesti NSV Teaduste Akadeemia Toimetised, Keemia, Geoloogia, 17, 409–424.
Mark-Kurik, E. & Põldvere, A. 2012. Devonian stratigraphy in Estonia: current state and problems. Estonian Journal of Earth Sciences, 61, 33–47.
Moloshnikov, S. V. 2009. Oredezhosteus, a new psammosteid genus (Heterostraci, Psammosteiformes) from the Lower Frasnian (Upper Devonian) of the Main Devonian Field. Paleontological Journal, 43, 197–200.
Novitskaya, L. I. 1965. Microstructure of some Psammosteida. In Psammosteidy (Agnatha, Psammosteidae) dеvоnа SSSR [Devonian Psammosteids (Agnatha, Psammosteidae) of the USSR] (Obruchev, D. V. & Mark-Kurik, E. Yu., eds), pp. 257–282. Institute of Geology, Academy of Sciences of the Estonian SSR, Tallinn [in Russian].
Novitskaya, L. I. 1971. On the diagnostic evaluation of the ornamentation of jawless vertebrates and fishes. Paleontologicheskij Zhurnal, 4, 82–96 [in Russian].
Novitskaya, L. I. 2004. Subclass Heterostraci. In Iskopaemye pozvonochnye Rossii i sopredel¢nykh stran. Beschelyustnye i drevnie ryby [Fossil Vertebrates of Russia and Adjacent Countries. Agnathans and Early Fishes] (Novitskaya, L. I. & Afanassieva, O. B., eds), pp. 69–207. GEOS, Moscow [in Russian].
Obruchev, D. V. 1933. Opisanie chetyrekh novykh vidov ryb Leningradskogo devona [Description of four new fish species from the Leningrad Devonian]. Materials of CNIGRI, Paleontology and Stratigraphy, 1, 12–15 [in Russian].
Obruchev, D. V. 1941. Ostatki Aspidosteus gen. nov. (Heterostraci) iz verkhnego devona r. Lovati [Remains of Aspidosteus gen. nov. (Heterostraci) from the Upper Devonian of River Lovat]. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR, 8(4), 7–22 [in Russian, with English summary].
Obruchev, D. V. 1943a. Psammosteidy (Psammosteiformes) devona SSSR i Shpitsbergena i evolyutsiya beschelyustnykh [Devonian Psammosteids (Psammosteiformes) of the USSR and Spitzbergen and the Evolution of Agnatha]. Palaeontological Institute of the Russian Academy of Sciences, Moscow, Manuscript of the DSc Thesis [in Russian].
Obruchev, D. V. 1943. Yoglinia n. g., latest pteraspid from the Middle Devonian of the Leningrad District. Doklady Akademii Nauk SSSR, 41, 41–43 [in Russian, with English summary].
Obruchev, D. V. 1947. O rode Psammosteus (Heterostraci) [About genus Psammosteus (Heterostraci)]. Doklady Akademii Nauk SSSR, 56, 517–520 [in Russian, with English summary].
Obruchev, D. V. 1964. The ramus Agnatha. Jawless vertebrates. In Osnovy paleontologii [Basics of Paleontology], Vol. 11 (Obruchev, D. V., ed.), pp. 34–116. Nauka, Moscow [in Russian].
Obruchev, D. V. & Mark-Kurik, E. Yu. 1965. Psammosteidy (Agnatha, Psammosteidae) devona SSSR [Devonian Psammosteids (Agnatha, Psammosteidae) of the USSR]. Institute of Geology, Academy of Sciences of the Estonian SSR, Tallinn, 304 pp. [in Russian, with English summary].
Obruchev, D. & Mark-Kurik, E. 1968. On the evolution of the psammosteids (Heterostraci). Eesti NSV Teaduste Akadeemia Toimetised, Keemia, Geoloogia, 17, 279–284.
Ørvig, T. 1961. Notes on some early representatives of the Drepanaspida (Pteraspidomorphi, Heterostraci). Arkiv för Zoologi, 12, 515–535.
Pernègre, V. N. & Elliott, D. K. 2008. Phylogeny of the Pteraspidiforms (Heterostraci), Silurian–Devonian jawless vertebrates. Zoologica Scripta, 37, 391–403.
http://dx.doi.org/10.1111/j.1463-6409.2008.00333.x
Preobrazhensky, I. A. 1911. O nekotorykh predstavitelyakh sem. Psammosteidae Ag. [On some representatives of the family Psammosteidae Ag.]. Protokoly obshchestva estestvoispytatelej Yur¢evskogo universiteta. Sitzungsberichte der Naturforscher–Gesellschaft bei der Universität Jurjew (Dorpat), 19(3, 4), 21–36 [in Russian].
Stensiö, E. A. 1927. The Downtonian and Devonian vertebrates of Spitsbergen. Part I. Family Cephalaspidae. Skrifter Svalbard Nordishavet, 12, 1–397.
Stensiö, E. A. 1958. Les Cyclostomes fossiles. In Traite de Zoologie, Vol. 13, Part 1 (Grassé, P. P., ed.), pp. 173–425. Masson, Paris.
Tarlo, L. B. 1962. The classification and evolution of the Heterostraci. Acta Palaeontologica Polonica, 7, 249–290.
Traquair, R. H. 1896. The extinct vertebrata animals of the Moray Firth Area. In A Vertebrate Fauna of the Moray Basin, Vol. 2 (Harvie-Brown, J. H. & Buckley, T. E., eds), pp. 235–285. Edinburgh.
Traquair, R. H. 1900. Report on fossil fishes collected by the Geological Survey of Scotland in the Silurian rocks of the South of Scotland. Transactions of the Royal Society of Edinburgh, 39, 827–864.
http://dx.doi.org/10.1017/S0080456800035237
Trautschold, H. 1880. Ueber Dendrodus und Coccosteus. Verhandlungen der Russisch-Kaiserlichen Mineralogischen Gesellschaft zu St. Petersburg, Zweite Serie, 15, 139–156.
Vorobyeva, E. I. & Lyarskaya, L. A. 1968. Ostatki kisteperykh i dvoyakodyshashchikh iz amatskikh sloev Latvii i ikh zahkoronenie [Remains of crossopterygians and dipnoans from the Amata Beds in Latvia and their burial conditions]. In Ocherki po filogenii i sistematike iskopaemykh ryb i beschelyustnykh [Essays on the Phylogeny and Systematics of Fossil Fishes and Agnathans] (Obruchev, D. V., ed.), pp. 71–86. Nauka, Moscow [in Russian].
Weber, V. N. 1924. Materialy k geologii yugo-vostochnoj chasti 40-go lista [Materials on the geology of the southeastern part of the 40th map sheet]. Izvestiya Geologicheskogo Komiteta, 40, 103–139 [in Russian].
Woodward, A. S. 1910. A Guide to the Fossil Reptiles, Amphibians, and Fishes in the Department of Geology and Palaeontology in the British Museum (Natural History). Printed by order of the Trustees, London, 129 pp.
© 2016 Authors. This is an Open Access article distributed under the terms and conditions of the Creative Commons Attribution 4.0 International License.