In memory of an outstanding palaeoichthyologist Elga Mark-Kurik
The range of diversity of psammosteids from the family Psammosteidae is still poorly known. Here a new species, Psammosteus ramosus sp. nov. Glinskiy, from the Amata Regional Stage of the Main Devonian Field is described. Its morphology, ornamentation, histology of exoskeletal plates, and micromeric elements are compared with those of other representatives of the family Psammosteidae. The comparison shows a close relationship of the new species with Psammosteus falcatus Obruchev, P. kiaeri Halstead Tarlo and P. pectinatus Obruchev, a group of species that is significantly different from other representatives of the genus Psammosteus and constitutes a separate evolutionary lineage. On the basis of morphological and histological features we here differentiate in the fields of tesserae of Psammosteidae the discrete micromeric elements of the ‘basic type’, known in Psammosteus bergi (Obruchev), P. levis Obruchev, P. livonicus Obruchev, P. maeandrinus Agassiz, P. megalopteryx (Trautschold), P. praecursor Obruchev and Karelosteus weberi Obruchev, and micromeric elements of the ‘progressive type’, known in Psammosteus falcatus, P. cf. kiaeri and P. ramosus sp. nov. Glinskiy.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. 1937. A classification of fish-like vertebrates. Izvestiya Akademii Nauk SSSR (Bulletin of the Academy of Sciences of the USSR), Otdelenie matematicheskikh i estestvennykh nauk, seriya biologicheskaya, 4, 1277–1280.
Blieck, A. 2016. Heterostracan vertebrates and the Great Eodevonian Biodiversification Event – an essay. Palaeobiodiversity and Palaeoenvironments, doi:10.1007/s12549-016-0260-1.
https://doi.org/10.1007/s12549-016-0260-1
Blieck, A. R. M., Karatajūtė-Talimaa, V. N. & Mark-Kurik, E. 2002. Upper Silurian and Devonian heterostracan pteraspidomorphs (Vertebrata) from Severnaya Zemlya (Russia): a preliminary report with biogeographical and biostratigraphical implications. Geodiversitas, 24, 805–820.
Blom, H., Carlsson, A. & Marshall, J. E. A. 2006. Vertebrate micro-remains from the Upper Devonian of East Greenland with comments of the Frasnian–Famennian boundary. Bulletin of the Geological Society of Denmark, 53, 39–46.
Bystrow, A. P. 1959. The microstructure of skeleton elements in some vertebrates from Lower Devonian deposits of the USSR. Acta Zoologica, 40, 59–83.
Delsate, D., Blieck, A. & Steemans, P. 2004. A psammosteid heterostracan (Vertebrata: Pteraspidomorphi) from the Emsian (Lower Devonian) of the Grand Duchi of Luxembourg. Geologica Belgica, 7, 21–26.
Dineley, D. L. & Loeffler, E. J. 1976. Ostracoderm faunas of the Delorme and associated Siluro-Devonian formations North West Territories Canada. Special Papers in Palaeontology, 18, 1–214.
Eichwald, E. 1844. O rybakh¢¢ pervobytnogo okeana v¢¢ okrestnostyakh¢¢ Pavlovska [About fishes from the primeval ocean in the Pavlovsk surroundings]. Otechestvennye Zapiski, 36(9), 1–22 [in Russian].
Elliott, D. K. & Mark-Kurik, E. 2005. A review of the lateral line sensory system in psammosteid heterostracans. Revista Brasileia de Paleontologia, 8, 99–108.
Glinskiy, V. N. 2013. Kompleksy srednedevonskikh psammosteidnykh beschelyustnykh vostochnoj chasti Glavnogo devonskogo polya [The assemblages of Middle Devonian psammosteid agnathans from the eastern part of Main Devonian Field]. Vestnik Saint-Petersburg University, Series 7, Geology, Geography, 4, 62–71 [in Russian, with English summary].
Glinskiy, V. N. 2014. Novye dannye po Karelosteus weberi Obruchev (Agnatha: Heterostraci) [New data on Karelosteus weberi Obruchev (Agnatha: Heterostraci)]. In Paleontologiya v muzejnoj praktike. Sbornik nauchnykh rabot [Palaeontology in the Museum Practice. Collection of the Scientific Articles] (Naugolnykh, S. V., ed.), pp. 22–25. Media-Grand, Moscow [in Russian, with English summary].
Glinskiy, V. N. & Ivanov, A. O. 2015. The assemblages of psammosteid agnathans from the Middle-Late Devonian of the Andoma Hill (Russia). In IGCP596-SDS Symposium: Climate Change and Biodiversity Patterns in the Mid-Palaeozoic (Mottequin, B., Denayer, J., Königshof, P., Prestianni, C. & Olive, S., eds), Strata (Travaux de Géologie Sédimentaire et Paléontologie), Série 1: Communications, 16, 57–59.
Glinskiy, V. N. & Mark-Kurik, E. 2016. Revision of Psammosteus livonicus Obruchev (Agnatha, Heterostraci) from the Devonian Amata Regional Stage of the NW of the East European Platform. Estonian Journal of Earth Sciences, 65, 1–18.
https://doi.org/10.3176/earth.2016.02
Goodrich, E. S. 1909. Vertebrata Сraniata (First fascile: cyclostomes and fishes). In A Treatise on Zoology. Part IX (Lankester, R., ed.), pp. XVI+518. Adam and Charles Black, London.
Gross, W. 1933. Die Fische des Baltischen Devons. Palaeontographica, 79, 1–74.
Gross, W. 1942. Die Fischfaunen des baltischen Devons und ihre biostratigraphische Bedeutung. Korrespondenzblatt des Naturforscher – Vereins zu Riga, 64, 373–436.
Gross, W. 1963. Drepanaspis gemuendenensis Schlüter Neuuntersuchung. Palaeontographica. Abt. A, 121(4–6), 133–155.
Halstead, L. B. 1987. Agnathan extinctions in the Devonian. Mémoires de la Société Géologique de France, 150, 7–11.
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, 1–168.
Halstead Tarlo, L. B. 1967a. Major faunal provinces in the Old Red Sandstone of the Northern Hemisphere. In International Symposium on the Devonian System, Calgary (Oswald, D. H., ed.), pp. 1231–1238. Alberta Society of Petroleum Geologists, Calgary.
Halstead Tarlo, L. B. 1967b. The tessellated pattern of the dermal armour in the Heterostraci. Journal of the Linnean Society of London, Zoology, 47, 45–54.
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. O., Lukševičs, E. V., Stinkulis, G. V., Tovmasyan, K. A., Zupinš, I. A. & Beznosov, P. A. 2006. Stratigrafiya devonskikh otlozhenij Andomskoj gory [Stratigraphy of the Devonian deposits of Andoma Hill]. In Problemy geologii i mineralogii [Problems of Geology and Mineralogy] (Pystin, A. M., ed.), pp. 385–396. Geoprint, Syktyvkar [in Russian].
Ivanov, A., Lukševičs, E. & Glinskiy, V. 2011. Vertebrate assemblages from the Givetian–Frasnian boundary beds of the Borschovo locality (Leningrad region, Russia). In The Eighth Baltic Stratigraphical Conference. Abstracts (Lukševičs, E., Stinkulis, Ģ. & Vasiļkova, J., eds), p. 29. University of Latvia, Riga.
Karatajūtė-Talimaa, V. 1998. Determination methods for the exoskeletal remains of Early Vertebrates. Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe, 1, 21–51.
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.
https://doi.org/10.1002/jmor.20370
Kiaer, J. 1915. Upper Devonian fish remains from Ellesmere Land with remarks on Drepanaspis. Report of the Second Norwegian Arctic Expedition in the “Fram” 1898–1902, 33, 1–72.
Kiaer, J. 1932. The Downtonian and Devonian vertebrates of Spitsbergen. IV. Suborder Cyathaspida. A preliminary report edited by A. Heintz. Skrifter om Svalbard og Ishavet, 52, 1–26.
Lankester, E. R. 1868. The fishes of the Old Red Sandstone of Britain. Part I. – The Cephalaspididae. Palaeontographical Society, 21, 1–33.
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 Palaeozoj Rossii: regional'naya stratigrafiya, paleontologiya, geo- i biosobytiya. Materialy III Vserossijskogo soveshchaniya [Palaeozoic 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. A. 1981. Pantsirnye ryby devona Pribaltiki. Asterolepididae [Аrmoured Fishes from Devonian of the Baltic States. Asterolepididae]. Zinatne, Riga, 153 pp. [in Russian, with English summary].
Mark-Kurik, E. 1993. Notes on the squamation in psammosteids. Modern Geology, 18, 107–114.
Mark-Kurik, E. 1999. Psammosteid microremains from the Middle Devonian (Givetian) of Estonia. Modern Geology, 24, 1–21.
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. 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 Devonian of Leningrad Province]. Materialy Tsentral'nogo Nauchno-Issledovatel'skogo Geologo-Razvedochnogo Instituta. Paleontologiya i Stratigraphiya, 1, 12–15 [in Russian, with English summary].
Obruchev, D. V. 1943. Yoglinia n. g., latest pteraspid from the Middle Devonian of the Leningrad District. Сomptes Rendus (Doklady) de l'Académie des Sciences de l'URSS, 41, 41–43.
Obruchev, D. V. 1945. Évolyutsiya Agnatha [Evolution of Agnatha]. Zoologicheskij Zhurnal, 24(5), 257–272 [in Russian].
Obruchev, D. V. 1947a. Tip Chordata. Hordovyе. Podtip Vertebrata. Pozvonochnye [Phylum Chordata. Chordates. Subphylum Vertebrata. Vertebrates]. In Atlas rukovodyashchikh form iskopaemykh faun SSSR. Tom III. Devonskaya sistema [Atlas of the Guide Forms of the Fossil Faunas of the USSR. Volume III. Devonian] (Nalivkin, D. V., ed.), pp. 191–206. State Editorial Office for Geological Literature, USSR Ministry of Geology, Moscow–Leningrad [in Russian].
Obruchev, D. V. 1947b. O rode Psammosteus (Heterostraci) [About genus Psammosteus (Heterostraci)]. Doklady Akademii Nauk SSSR, 56, 517–520 [in Russian].
Obruchev, D. V. 1964. Vetv' Agnatha. Beschelyustnye [The ramus Agnatha. Jawless vertebrates]. In Osnovy paleontologii. Beschelyustnye, ryby [Basics of Palaeontology. Jawless Vertebrates, Fishes], 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].
Ørvig, T. 1951. Histologic studies of placoderms and fossil elasmobranchs. I: The endoskeleton, with remarks on the hard tissues of lower vertebrates in general. Arkiv för Zoologi, 2, 321–454.
Ørvig, T. 1977. A survey of odontodes (‘dermal teeth’) from developmental, structural, functional and phyletic points of view. In Problems in Vertebrate Evolution (Andrews, S. M., Miles, R. S. & Walker, A. D., eds), Linnean Society Symposium Series, 4, 53–75.
Rohon, J. V. 1901. Beiträge zur Anatomie und Histologie der Psammosteiden. Vĕstník Královské České Společnosti Nauk, 16, 1–31.
Schlüter, С. A. J. 1887. Über Panzerfische und legte neue Arten aus dem rheinisch-westfalischen Devon vor. Verhandlungen des Naturhistorischen Vereines der Preussischen Rheinlande und Westphalens (Naturwissenschaftliche Sektion), 44, 120–128.
Selden, P. A. 2003. A new tool for fossil preparation. The Geological Curator, 7, 337–339.
Tarlo, L. B. 1961. Psammosteids from the Middle and Upper Devonian of Scotland. The Quarterly Journal of the Geological Society of London, 117, 193–212.
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. 1897. Additional notes on the fossil fishes of the Upper Old Red Sandstone of the Moray Firth Area. Proceedings of the Royal Physical Society of Edinburgh, 13, 376–385.
Trautschold, H. 1880. Ueber Dendrodus und Coccosteus. Verhandlungen der Russisch-Kaiserlichen Mineralogischen Gesellschaft zu St. Petersburg, Zweite Serie, 15, 139–156.
Vaškaninová, V. & Kraft, P. 2016. A unique occurrence of a psammosteid heterostracan on the peri-Gondwanan shelf in the Lower/Middle Devonian boundary marine deposits. Fossil Imprint, 72, 155–160.