Phylogenetic relationships of psammosteid heterostracans (Pteraspidiformes), Devonian jawless vertebrates

  • Vadim Glinskiy Institute of Earth Sciences, Saint Petersburg State University, Universitetskaya nab., 7–9, Saint Petersburg, 199034, Russian Federation http://orcid.org/0000-0001-9151-1161

Abstract

Psammosteid heterostracans are a group (suborder Psammosteoidei) of Devonian-age jawless vertebrates, which is included in the order Pteraspidiformes. The whole group of psammosteids is represented by numerous species (more than 40); their phylogenetic relationships are still poorly known and deserve further study. Classical researchers of the psammosteids, such as D. Obruchev, E. Mark-Kurik and L. Halstead Tarlo, had different views on the phylogeny of the group (e.g. origins and evolution of Psammosteus). To check the modern hypotheses of psammosteid origins from various Pteraspidiformes and to clarify psammosteid interrelationships, the most complete phylogeny of this group (38 ingroup taxa + juvenile Drepanapsis) is presented here. Different methods of data analysis were used to explore the psammosteid data set, including equally weighted characters versus implied weighting. According to the results of the phylogenetic analysis, the monophyletic status of the group and their early development from the Pteraspidiformes are supported. The diagnoses and interrelationships of many taxa are clarified. Two new genera are proposed (Vladimirolepis gen. nov. and Elgaia gen. nov.), and two subfamilies are erected (Placosteinae subfam nov. and Psammosteinae Traquair).

Keywords:

Agnatha, Heterostraci, Pteraspidiformes, psammosteids, phylogeny, classification

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References

Agassiz, L. 1833–1843. Recherches sur les Poissons fossiles. Tableau général des Poissons fossiles, rangés par terrains, vol. 1. Neuchâtel.

Agassiz, L. 1844. Monographie des Poissons fossiles du vieux grè s rouge ou système Dévonien (Old Red Sandstone) des Isles Britanniques et de Russie. Neuchâtel: Jent & Gaßmann (Solothurn).

Agassiz, L. 1845. Lettres sur les Poissons fossiles du système Dévonien de la Russie; pp. 397–418 in Murchison, R. I., dе Vеrnеuil, Е. & A. de Кеуsеrling, C. А. Géologie de la Russie d’Europe et des montagnes de l’Oural, vol. 2. Paris.

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:1277–1280.

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 5(2):87–517.

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 20:1–289.

Blieck, A. 1984. Les Hétérostracés Ptéraspidiformes, agnathes du Silurien-Dévonien du continent Nord-Atlantique et des blocs avoisinants: revision systématique, phylogénie, biostratigraphie, biogéographie. Paris: Cahiers de Paléontologie (section Vertébrés), éditions du CNRS.

Blieck, A. 1991. Reappraisal of the heterostracans (Agnathan vertebrates) of Northern Ireland. Irish Journal of Earth Sciences 11:65–69.

Blieck, A. R. M., Elliott, D. K., and Gagnier, P.-Y. 1991. Some questions concerning the phylogenetic relationships of heterostracans, Ordovician to Devonian jawless vertebrates; pp. 1–17 in C. Mee-Mann, L. Yu.-hai and G. Zhang (eds), Early vertebrates and related problems of evolutionary biology. Beijing: Science Press.

Brotzen, F. 1936. Beiträge zur Vertebratenfauna des westpodolischen Silurs und Devons. I. Protaspis arnelli n. sp. und Brachipteraspis n. gen. latissima Zych. Arkiv för Zoologi 28A (22):1–52.

Bystrow, A. P. 1955. Mikrostruktura pantsyrya beschelyustnykh pozvonochnykh silura i devona [Microstructure of the armor of the jawless vertebrates from Silurian and Devonian]; pp. 472–523 in E. M. Murzaev (ed.), Pamyati akademika L. S. Berga. Sbornik rabot po geografii i biologii. Publishing of AN USSR, Moscow-Leningrad.

Bystrow, A. P. 1959. The microstructure of skeleton elements in some vertebrates from Lower Devonian deposits of the USSR. Acta Zoologica 40:59–83. https://doi.org/10.1111/j.1463-6395.1959.tb00392.x

Denison, R. H. 1968. Middle Devonian fishes from the Lemhi Range of Idaho. Fieldiana Geology 16:269–287.

Doss, B. 1915. Ein Fund von Psammosteus arenatus Ag. bei Wenden in Livland. Korrespondenzblatt des Naturforscher-Vereins zu Riga 57:77–78.

Eichwald, E. 1860. Lethaea Rossica, ou Paléontologie de la Russie, décrite et figurée. Premier Volume. Ancienne periode. Stuttgart: E. Schweizerbart.

Elliott, D. K. 1984. A new subfamily of the Pteraspididae (Agnatha, Heterostraci) from the Upper Silurian and Lower Devonian of Arctic Canada. Palaeontology 27:169–197.

Elliott, D. K., and Mark-Kurik, E. 2005. A review of the lateral line sensory system in psammasteid heterostracans. Revista Brasileia de Paleontologia 8:99–108.

Elliott, D., Mark-Kurik, E., and Daeschler, E. B. 2004. A revision of Obruchevia (Psammosteida: Heterostraci) and the description of a new obrucheviid from the Late Devonian of the Canadian Arctic. Acta Universitatis Latviensis 679:22–45.

Glinskiy, V. N. 2014. New records of psammosteids (Heterostraci) from the Aruküla Regional Stage (Middle Devonian) of the Leningrad Region, Russia. Paleontological Journal 48:980–991. https://doi.org/10.1134/S0031030114090032

Glinskiy, V. 2017. Evolutionary lineages in the family Psammosteidae (Agnatha: Pteraspidiformes): pp. 39–40, in Ginter, M. (ed.), 14th International Symposium on Early and Lower Vertebrates (Cheçiny, Poland, 2017). Ichthyolith Issues, Special Publication, 13.

Glinskiy, V. 2018. Resul’taty analiza filogeneticheskikh svjazej devonskykh psammosteidnykh beschelyustnykh (Pteraspidiformes: Psammosteida). [Results of analysis of the phylogenetic relationships of the Devonian psammosteid agnathans (Pteraspidiformes: Psammosteida)]: pp. 189–190, in Bogdanova, T. M. et al. (eds), Fundamental'naya i prikladnaya paleontologija. Materialy LXIV sessii Paleontologicheskogo obshestva (Saint Petersburg, Russia, 2–6 April, 2018).

Glinskiy, V. N., and 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

Glinskiy, V. N., and Nilov, S. P. 2017. A new psammosteid (Agnatha, Heterostraci) from the Amata Regional Stage of the Main Devonian Field and morpho-histological types of discrete micromeric elements in the family Psammosteidae. Estonian Journal of Earth Sciences 66:59–76. https://doi.org/10.3176/earth.2017.05

Glinskiy, V. N., and Pinakhina, D. V. 2018. New data on psammosteid heterostracans (Pteraspidomorpha) and acanthodians (Acanthodii) from the Pärnu Regional Stage (Lower Eifelian, Middle Devonian) of Estonia. Estonian Journal of Earth Sciences 67:76–87. https://doi.org/10.3176/earth.2018.05

Goloboff, P. A. 2014. Extended implied weighting. Cladistics 30: 260–272. https://doi.org/10.1111/cla.12047

Goloboff, P. A., Farris, J. S. and Nixon, K. C. 2008. Cladistics TNT, a free programm for the phylogenetic analysis. Cladistics 24: 774–786. https://doi.org/10.1111/j.1096-0031.2008.00217.x

Gross, W. 1930. Die Fische des mittleren Old Red Süd-Livlands. Geologische und Palaeontologische Abhandlungen 22 (N. F. 18):123–156.

Gross, W. 1933a. Die Fische des Baltischen Devons. Palaeontographica 79A:1–74.

Gross, W. 1933b. Die phylogenetische Bedeutung der altpaläozoischen Agnathen und Fische. Paläontologische Zeitschrift 15:102–137.

Gross, W. 1935. Histologische Studien am Aussenskelett fossiler Agnathen und Fische. Palaeontographica 83А:1–60.

Gross, W. 1942. Die Fischfaunen des baltischen Devons und ihre biostratigraphische Bedeutung. Korrespondenzblatt des Naturforscher-Vereins zu Riga 64:373–436.

Gross, W. 1937a. Die Wirbeltiere des rheinischen Devons. Teil I. Abhandlungen der Preußischen Geologischen Landesanstalt Neue Folge:1–81.

Gross, W. 1937b. Die Wirbeltiere des rheinischen Devons. Teil II. Abhandlungen der Preußischen Geologischen Landesanstalt Neue Folge:1–83.

Gross, W. 1963. Drepanaspis gemuendenensis Schlüter. Neuuntersuchung. Palaeontographica. Abteilung A 121 (4–6):133–155.

Gross, W. 1967. Bemerkungen zum System und zur Phylogenie der Agnathen und Fische. Colloques Internationaux du Centre National de la Recherche Scientifique. Problèmes actuels de paléontologie (Évolution des vertébrés) 163:73–79.

Halstead, L. B. 1969. Calcified tissues in the earliest vertebrates. Calcified Tissue Research 3:107–124. https://doi.org/10.1007/BF02058654

Halstead, L. B. 1973. The heterostracan fishes. Biological reviews of the Cambridge Philosophical Society 48:279–332. https://doi.org/10.1111/j.1469-185X.1973.tb01005.x

Halstead, L. B. 1993. Agnatha; pp. 573–581 in M. J. Benton. The Fossil Record 2. London: Chapmann & Hall.

Halstead Tarlo, L. B. 1964a. 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. 1964b. The origin of bone; pp. 3–17 in H. J. J. Blackwood, ed. Bone and Tooth: Proceedings of the First European Symposium held at Somerville College, Oxford, April 1963. Oxford: A Pergamon Press Book.

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.

Ilyes, R. R., and Elliott, D. K. 1994. New Early Devonian pteraspidids (Agnatha, Heterostraci) from East-Central Nevada. Journal of Paleontology 68:878–892. https://doi.org/10.1017/S0022336000026330

Janvier, P. 1996. Early Vertebrates. Oxford and New York: Clarendon Press (Oxford University Press).

Johanson, Z., Smith, M., Kearsley, A., Pilecki, P., Mark-Kurik, E., and 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 9:1–5. http://doi.org/10.1098/rsbl.2013.0144

Keating, J. N., Marquart, C. L., and 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. 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.

Lebedev, O. A., Mark-Kurik, E. Y., Karatajūtė-Talimaa, V. N., Lukševičs, E. V., and Ivanov, A. О. 2009. Bite marks as evidence of predation in early vertebrates. Acta Zoologica 90 (Suppl. 1):344–356. https://doi.org/10.1111/j.1463-6395.2008.00344.x

Lyarskaya, L. A. 1971. Novye nakhodki ostatkov psammosteid iz shvyantojskogo gorizonta Latvii [New finds of psammosteid remains from the Šventoji Formation of Latvia]; pp. 97–104 in A. A. Grigelis (ed.), Paleontologiya i stratigraphiya Pribaltiki i Belorussii. Sbornik 3. Mintis, Vilnius.

Mark, E. Y. 1955. Psammosteidy (Agnatha) tartuskogo i gaujskogo gorizontov devona Estonskoj SSR [Psammosteids (Agnatha) from Tartu and Gauja formations of Devonian deposits of Estonian SSR], Tallinn.

Mark-Kurik, E. Y. 1966. O nekotorykh povrezhdeniiakh ekzoskeleta psammosteid (Agnatha). [On the some injuries of the exoskeleton of the psammosteids (Agnatha)]; pp. 55–60 in R. G. W. Hecker (ed.), Organism i sreda v geologicheskom proshlom. Nauka, Moskow.

Mark-Kurik, E. 1968. New finds of psammosteids (Heterostraci) in the Devonian of Estonia and Latvia. Proceedings of the Estonian Academy of Sciences. Chemistry & Geology 17:409–424.

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.

Moloshnikov, S. V. 2001. New data on Pycnosteus palaeformis Preobrazhensky (Heterostraci, Psammosteiformes) from Aruküla deposits of Leningrad Region. Paleontological Journal 35(4):410–414.

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(2):197–200. https://doi.org/10.1134/S0031030109020117

Novitskaya, L. I. 1965. Mikrostroenie nekotorykh Psammosteida [Microstructure of some Psammosteida]; pp. 257–282 in D. V. Obruchev, E. Y. Mark-Kurik, Psammosteidy (Agnatha, Psammosteidae) dеvоnа SSSR [Devonian psammosteids (Agnatha, Psammosteidae) of the USSR]. Tallinn: Institute of Geology, Academy of Sciences of the Estonian SSR.

Novitskaya, L. I. 2004. Podklass Heterostraci [Suborder Heterostraci]; pp. 69–207 in L. I. Novitskaya, (ed.), Fossil vertebrates of Russia and adjacent countries. Agnathans and early fishes. GEOS, Moscow.

Obruchev, D. V. 1940. O nekotorykh psammosteidakh Leningradskogo i Pribaltijskogo srednego devona. [On some psammosteids from the Devonian of Leningrad district and Baltic States]. Doklady Akademii Nauk SSSR 28(8):766–768.

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 Lovat’ River]. Trudy Paleontologicheskogo Instituta 8(4):7–22.

Obruchev, D. V. 1943a. 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(1):41–43.

Obruchev, D. V. 1943b. Novaya rekonstrukcija Drepanaspis [The new reconstruction of Drepanaspis]. Doklady Akademii Nauk SSSR 43(6):269–272.

Obruchev, D. V. 1943c. Psammosteidae devona SSSR i Spitsbergena i evolyutsiya Agnatha [Psammosteidae from the Devonian of USSR and Spitsbergen and evolution of Agnatha]. Biological Department of the Academy of Sciences of USSR. Moskow.

Obruchev, D. V. 1945. Evolutsija Agnatha [The Evolution of Agnatha] Zoological Zhournal 24:257–271.

Obruchev, D. V. 1947a. Tip Chordata. Khordovye. Podtip Vertebrata. Pozvonochnye [Phylum Chordata. Chordates. Subphylum Vertebrata. Vertebrates]; pp. 191-206 in D. Nalivkin, (ed.), Atlas of the guide forms of the fossil faunas of the USSR. Volume III. Devonien, vol. 3. State Editorial Office for Geological Literature USSR Ministry of Geology, Moskow-Leningrad.

Obruchev, D. V. 1947b. O rode Psammosteus (Heterostraci) [About genus Psammosteus (Heterostraci)]. Doklady Akademii Nauk SSSR 56(5):517–520.

Obruchev, D. V. 1958. K biostratigrafii ikhtiofaun nizhnego i srednego paleozoya SSSR [To the biostratigraphy of the ichthyofauna of the Lower and Middle Paleozoic USSR]. Sovetskaya Geologiya 11:40–53.

Obruchev, D. V. 1961. Rod Tartuosteus (Psammosteidae) iz srednego devona Pribaltiki [Genus Tartuosteus (Psammosteidae) from the Middle Devonian of Baltic States]. Paleontological Journal 2:107–111.

Obruchev, D. V. 1964. Vetv’ Agnatha. Beschelyustnye [The ramus Agnatha. Jawless vertebrates]; pp. 34–116 in D. V. Obruchev (ed.), Osnovy paleontologii. Beschelyustnye, ryby. Basics of Palaeontology. Jawless Vertebrates, Fisches, vol. 11. Nauka, Moscow.

Obruchev, D. V. 1967. On the evolution of the Heterostraci. Colloques Internationaux du Centre National de la Recherche Scientifique. Problèmes actuels de paléontologie (Évolution des vertébrés) 163:37–43.

Obruchev, D. V. 1968. Ob evolyutsii Heterostraci [On the evolution of Heterostraci]; pp. 21–23 in D. V. Obruchev, (ed.), Essays on phylogeny and taxonomy of fossil fishes and agnathans. Nauka, Moscow.

Obruchev, D. V. 1972. Nekotorye kriterii filogeneticheskikh issledovanij na primere nizshikh pozvonochnykh [Some criteria of phylogenetic studies on the example of lower vertebrates]. Paleontological Journal 3:56–70.

Obruchev, D. V., and Mark-Kurik, E. Yu. 1965. Devonian psammosteids (Agnatha, Psammosteidae) of the USSR. Tallinn: Institute of Geology, Academy of Sciences of the Estonian SSR.

Obruchev, D. V., and Mark-Kurik, E. 1968. On the evolution of the psammosteids (Heterostraci). Proceedings of the Academy of Sciences of the Estonian SSR, Chemistry & Geology 17:279–284.

Ørvig, T. 1961. Notes on some early representatives of the Drepanaspida (Pteraspidomorphi, Heterostraci). Arkiv för Zoologi 2(12):515–535.

Pernègre, V. N. 2002. The genus Doryaspis White (Heterostraci) from the Lower Devonian of Vestspitsbergen, Svalbard. Journal of Vertebrate Paleontology 22(4):735–746. https://doi.org/10.1671/0272-4634(2002)022[0735:TGDWHF]2.0.CO;2

Pernègre, V. N., and Elliott, D. K. 2008. Phylogeny of the Pteraspidiformes (Heterostraci), Silurian–Devonian jawless vertebrates. Zoologica Scripta 37(4):391–403. https://doi.org/10.1111/j.1463-6409.2008.00333.x

Pernègre, V., and Goujet, D. 2007. The genus Gigantaspis Heintz, 1962 (Vertebrata, Heterostraci) from the Lower Devonian of Spitsbergen. Palaeontology 50(2):323–346. https://doi.org/10.1111/j.1475-4983.2007.00638.x

Plax, D. P. 2010. Rannefrnskaya ikhitofauna severa Belarusi [Lower Frasnian ichthyofaunal from the North of Belarus]. Litasfera 1(32):60–81.

Randle, E., and Sansom, R. S. 2016. Exploring phylogenetic relationships of Pteraspidiformes heterostracans (stemgnathostomes) using continuous and discrete characters. Journal of Systematic Palaeontology 14:1–17. https://doi.org/10.1080/14772019.2016.1208293

Randle, E., and Sansom, R. S. 2017. Phylogenetic relationships of the ‘higher heterostracans’ (Heterostraci: Pteraspidiformes and Cyathaspididae), extinct jawless vertebrates. Zoological Journal of the Linnean Society, 14:1–17. https://doi.org/10.1093/zoolinnean/zlx025

Růžička, R. 1929. Psammosteus (Ganosteus) perneri n. sp. Příspěvek ku poznáni ryb českého paleozoika. Vestnik Státního Geologického Ústavu Čsl. Republiky 5:1–7.

Stensiö, E. A. 1927. The Downtonian and Devonian vertebrates of Spitsbergen. Part I. Family Cephalaspidae. Skrifter Svalbard Nordishavet 12:1–391.

Stensiö, E. A. 1958. Les cyclostomes fossiles ou ostracodermes; pp. 173–425 in P.-P. Grassé, (ed.), Traité de Zoologie, vol. 13:1. Masson, Paris.

Tarlo, L. B. 1957. A preliminary note on new ostracoderms from the Lower Devonian (Emsian) of Central Poland. Acta Palaeontologica Polonica 11:225–233.

Tarlo, L. B. 1961. Psammosteids from the Middle and Upper Devonian of Scotland. The Quarterly Journal of the Geological Society of London 117:193–213. https://doi.org/10.1144/gsjgs.117.1.0193

Tarlo, L. B. 1962. The classification and evolution of the Heterostraci. Acta Palaeontologica Polonica 7:249–290.

Tarlo, L. B. H. 1967. Agnatha (Part II: Documentation of the fossil record); pp. 629–636 in W. B. Harland et al. (eds), The Fossil Record. Geological Society of London, London.

Traquair, R. H. 1896. The extinct vertebrate animals of the Moray Firth Area; pp. 235–285 in Harvie-Brown, J. A. and Buckley, T. E. (eds), A vertebrate fauna of the Moray Basin. Vol. 2. Edinburgh: David Douglas.

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.

Traquair, R. H. 1899. 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. https://doi.org/10.1017/S0080456800035237

Traquair, R. H. 1900. Notes on Drepanaspis gemündenensis, Schlüter. The Geological Magazine 4(7):153–159. https://doi.org/10.1017/S0016756800177027

Vaškaninová, V., and 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(3–4):155–160. https://doi.org/10.14446/FI.2016.155

Westoll, T. S. 1967. Radotina and other tesserate fishes. Zoological Journal of the Linnean Society 47:83–98. https://doi.org/10.1111/j.1096-3642.1967.tb01397.x

White, E. I. 1935. The Ostracoderm Pteraspis Kner and the relationships of the agnathous vertebrates. Philosophical Transactions of the Royal Society B 225:381–457. https://doi.org/10.1098/rstb.1935.0015

Woodward, A. S. 1891. Catalogue of the fossil fishes in the British Museum (Natural History). Pt. 2. London: Printed by order of the Trustees.

Woodward, A. S. 1910. A guide to the fossil reptiles, amphibians, and fishes in the Departement of Geology and Palaeontology in the British Museum (Natural History). 9-th edition. London, British Museum.
Published
2018-05-04
How to Cite
Glinskiy, V. (2018). Phylogenetic relationships of psammosteid heterostracans (Pteraspidiformes), Devonian jawless vertebrates. Biological Communications, 62(4), 219–243. https://doi.org/10.21638/11701/spbu03.2017.402
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