Bryozoan fauna from the Kunda Stage (Darriwilian, Middle Ordovician) of Estonia and NW Russia


Authors: Ernst A

Published in: Bulletin of Geosciences, volume 97, issue 1; pages: 33 - 68; Received 30 September 2021; Accepted in revised form 20 December 2021; Online 23 January 2022

Keywords: Ordovician; Darriwilian; Bryozoa; systematics; palaeogeography; palaeoecology,

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Bryozoan fauna from several outcrops of sediments of the Kunda Stage (Darriwilian, Middle Ordovician) of Estonia and NW Russia contains 18 species which belong to the palaeostomate orders Cystoporata, Esthonioporata, Trepostomata, and Cryptostomata. The studied material comes mainly from the collection of the Department of Geology, Tallinn University of Technology, as well as was specimens I have collected. One genus with one species and one more species are new: Pakripora cavernosa gen. et sp. nov. and Dianulites pakriensis sp. nov. Four bryozoans are identified to genus level only. One cryptostome bryozoan shows unique morphology unknown in any similar genera. In general, the studied fauna shows close relations to other localities within the Baltic region, with some few relations to the Ordovician of China. The bryozoans developed massive and erect colonies, showing adaptation to the shallow to moderately deep environment with moderate water energy and relatively high level of sedimentation.


Amini Z.Z., Adabi M.H., Burrett C.F. & Quilty P.G. 2004. Bryozoan distribution and growth form associations as a tool in environmental interpretation, Tasmania, Australia. Sedimentary Geology 167(1–2), 1–15. View article

Anstey, R.L. & Perry, T.G. 1970. Biometric procedures in taxonomic studies of Paleozoic bryozoans. Journal of Paleontology 44, 383–398.

Astrova, G.G. 1964. A new order of the Paleozoic Bryozoa. Paleontologicheskiy Zhurnal 1964(2), 22–31. [in Russian]

Astrova, G.G. 1965. Morphology, history of development and system of the Ordovician and Silurian Bryozoa. Trudy Paleontologicheskogo Instituta 106, 1–432. [in Russian]

Astrova, G.G. 1978. The history of development, system, and phylogeny of the Bryozoa: Order Trepostomata. Trudy Paleontologicheskogo Instituta 169, 1–240. [in Russian]

Astrova, G.G. & Morozova, I.P. 1956. On systematics of the bryozoans of the Order Cryptostomata. Doklady Akademii Nauk SSSR 110(4), 661–664. [in Russian]

Banta, W.C., McKinney, F.K. & Zimmer, R.L. 1974. Bryozoan monticules: excurrent water outlets? Science 185, 783–784. View article

Bassler, R.S. 1911. The early Paleozoic Bryozoa of the Baltic Provinces. Bulletin of the Smithsonian Institution, United States National Museum 77, 1–382.

Bassler, R.S. 1913. Bryozoa, 314–355. In Zittel, K.A. & Eastman, C.R (eds) Text-book of Paleontology, Vol. 1. Macmillan & Co., London.

Bassler, R.S. 1936. Nomenclatorial notes on fossil and Recent Bryozoa. Journal of the Washington Academy of Science 26, 156–162.

Bassler, R.S. 1952. Taxonomic notes on genera of fossil and Recent Bryozoa. Journal of the Washington Academy of Sciences 42, 381–385.

Borg, F. 1926. Studies on Recent cyclostomatous Bryozoa. Zoologiska Bidrag fran Uppsala 10, 181–507.

Brood, K. 1984. Bryozoan ecology in the Silurian of Gotland. Special Papers in Palaeontology 32, 211–224. View article

Buttler, C. & Massa, D. 1996. Late Ordovician bryozoans from carbonate buildups, Tripolitania, Libya, 63–68. In Gordon, D.P., Smith, A.M. & Grant-Mackie, J.A. (eds) Bryozoans in Space and Time. Proceedings of the 10th International Bryozology Conference, Wellington, New Zealand, 1995. National Institute of Water and Atmospheric Research Ltd, Wellington.

Buttler, C., Cherns, L. & Massa, D. 2007. Bryozoan mud-mounds from the Upper Ordovician Jifarah (Djeffara) Formation of Tripolitania, north-west Libya. Palaeontology 50(2), 479–494. View article

Cocks, R.L.M. & Torsvik, T.H. 2005. Baltica from the late Precambrian to mid-Palaeozoic times: the gain and loss of a terrane’s identity. Earth-Science Reviews 72, 39–66. View article

Conti, S. & Serpagli, E. 1987. Functional morphology of the cap-like apparatus in autozooids of a Palaeozoic trepostome bryozoan. Lethaia 20, 1–20. View article

Cook, P.L. 1977. Colony-wide water currents in living Bryozoa. Cahiers de Biologie Marine 18, 31–47.

Dana, J.D. 1846. Zoophytes. United States Exploring Expedition 7, 1–740. View article

Dronov, A. & Rozhnov, S. 2007. Climatic changes in the Baltoscandian basin during the Ordovician: sedimentological and palaeontological aspects. Acta Geologica Sinica 46, 108–113.

Dybowski, W. 1877. Die Chaetetiden der ostbaltischen Silur-Formation. Russische Kaiserliche Mineralogische Gesellschaft zu St. Petersburg, Verhandlungen, Series 2(14), 1–134.

Ehrenberg, C.G. 1831. Symbolae Physicae, seu Icones et descptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegiptum, Nubiam, Dongalaam, Syriam, Arabiam et Habessiniam, studia annis 1820–25, redirent. Pars Zoologica, 4, Animalia Evertebrata exclusis Insectis. 10 pls. Berolini.

Eichwald, E. 1829. Zoologia specialis quam expositis animalibus tum vivis, fom fossilibus potissimum Rossiae in universum, et Poloniae in specie, vol. 1. Bryozoa. 314 pp. Typis Josephi Zawadzki, Vilnae. View article

Eichwald, E. 1856. Beitrag zur geographischen Verbreitung der fossil Thiere Russlands. Bulletin de la Société des Naturelles de Moscovian 29, 91–96, 448–466.

Eichwald, E. 1860. Lethaea Rossica, ou Paléontologie de la Russie. I. Ancienne Periode. 1657 pp. Schweizerbart, Stuttgart.

Ernst, A. 2018. Diversity dynamics of Ordovician Bryozoa. Lethaia 51, 198–206. View article

Ernst, A. & Nakrem, H.A. 2011. Stenolaemate bryozoans from the Mjosa Formation (Late Ordovician, Katian) of Helgoya (Bergevika), southern Norway. Norwegian Journal of Geology 91, 163–180.

Ernst, A. & Nakrem, H.A. 2012. Late Ordovician (Sandbian) bryozoans and their depositional environment, Furuberget Formation, Mjosa District, Oslo Region, Norway. Bulletin of Geosciences 87(1), 21–44. View article

Fedorov, P.V., Koromyslova, A.V. & Martha, S.O. 2017. The oldest bryozoans of Baltoscandia from the lowermost Floian (Ordovician) of north-western Russia: two new rare, small and simple species of Revalotrypidae. Paläontologische Zeitschrift 91(3), 353–363. View article

Gorjunova, R.V. 1988. On the systematic position of the genus Revalotrypa. Paleontologicheskii Zhurnal 1988(2), 31–36. [in Russian]

Gorjunova, R.V. 1996. Phylogeny of the Paleozoic Bryozoa. Trudy Paleontologischeskogo Instituta 267, 1–161.

Gorjunova, R.V. 2005. Ralfimartitidae, a new family of Paleozoic bryozoans of the order Trepostomida. Paleontologicheskii Zhurnal 2, 51–63. [in Russian]

Gorjunova, R.V. 2009. Evolution of the colonial growth habit in the Ordovician bryozoans of the Class Stenolaemata: feeding adaptation (Leningrad Region, Russia). Paleontologicheskii Zhurnal 43 (Supplement 11), 1390–1408. View article

Gorjunova, R.V. & Koromyslova, A.V. 2008. A new genus of the Arenigian bryozoans, Lynnopora, and its systematic position in the family Revalotrypidae Gorjunova, 1988. Paleontological Journal 42(5), 491–499. View article

Gorjunova, R.V. & Lavrentjeva, V.D. 1987. New genus Prophyllodictya, and the most ancient representative of cryptostome Bryozoa. Paleontologicheskii Zhurnal 1987(1), 41–51. [in Russian]

Gorjunova, R.V. & Lavrentjeva, V.D. 1993. Morphology and system of the cryptostome bryozoans. Trudy Paleontologischeskogo Instituta 257, 1–150. [in Russian]

Hageman, S.J. 1993. Effects of nonnormality on studies of the morphological variation of a rhabdomesine bryozoan, Streblotrypa (Streblascopora) prisca (Gabb and Horn). The University of Kansas Paleontological Contributions 4, 1–13. View article

Hageman, S.J., Bone, Y., McGowran, B. & James, N.P. 1997. Bryozoan colonial growth forms as paleoenvironmental indicators: Evaluation of methodology. Palaios 12, 406–419. View article

Hall, J. 1852. Palaeontology of New York, Volume 2, Organic Remains of the Lower Middle Division of the New York System. New York Geological Survey, Natural History of New York v. 6, 40–52, 144–173.

Hansen, T., Nielsen, A.T. & Bruton, D.L. 2011. Palaeoecology in a mud-dominated epicontinental sea: A case study of the Ordovician Elnes Formation, southern Norway. Palaeogeography, Palaeoclimatology, Palaeoecology 299, 348–362. View article

Hints, O., Ainsaar, L., Männik, P. & Meidla, T. (eds) 2008. The Seventh Baltic Stratigraphical Conference. Abstracts and Field Guide. 46 pp. Geological Society of Estonia, Tallinn.

Kaljo, D. & Nestor, H. 1990. Field meeting Estonia, 1990: An Excursion Guidebook. 209 pp. Institute of Geology, Estonian Academy of Sciences, Tallinn.

Key, M.M., Jr 1990. A new family of trepostome bryozoans from the Ordovician Simpson Group of Oklahoma. Journal of Paleontology 64, 700–724. View article

Key, M.M., Jr & Judd, A.B. 1994. Phylogenetic relationships of the Middle Ordovician trepostome bryozoans Sonninopora and Bimuropora. Journal of Paleontology 68, 241–257. View article

Koromyslova, A.V. 2004. Bryozoans of the Genus Diplotrypa Nicholson, 1879 from the Middle Ordovician of the Leningrad Region. Paleontologicheskii Zhurnal 2004(6), 47–49. [in Russian]

Koromyslova, A.V. 2007. Morphogenesis, phylogenetic relationships, and distribution of the bryozoans of the genus Orbipora Eichwald, 1856 (Order Trepostomida). Paleontological Journal 41(2), 132–145. [in Russian] View article

Koromyslova, A.V. 2011. Bryozoans of the Latorp and Volkhov Horizons (Lower-Middle Ordovician) of the Leningrad Region. Paleontological Journal 45(8), 887–980. View article

Koromyslova, A. & Fedorov, P. 2021. The oldest bifoliate cystoporate and two other bryozoan taxa from the Dapingian (Middle Ordovician) of north-western Russia. Journal of Paleontology 95(1), 24–39. DO 10.1017/jpa.2020.73View article

Lavrentjeva, V.D. 1975. A new bryozoan genus of the family Phylloporinidae. Paleontologicheskii Zhurnal 1975(4), 138–140. [in Russian]

Lavrentjeva, V.D. 1985. Bryozoans of the sub-order Phylloporinida. Trudy Paleontologicheskogo Instituta 214, 1–100. [in Russian]

Lidgard, S. 1981. Water flow, feeding, and colony form in an encrusting cheilostome, 135–142. In Larwood, G.P. & Nielsen, C. (eds) Recent and Fossil Bryozoa. Olsen & Olsen, Fredensborg.

Ma, J.-Y., Buttler, C.J. & Taylor, P.D. 2014. Cladistic analysis of the ‘trepostome’ Suborder Esthonioporina and the systematics of Palaeozoic bryozoans, 153–161. In Rosso, A., Wyse Jackson, P.N. & Porter, J.S. (eds) Bryozoan Studies 2013. Studi Trentini di Scienze Naturali 94.

Ma, J., Taylor, P.D., Xia, F. & Zhan, R. 2015. The oldest known bryozoan: Prophyllodictya (Cryptostomata) from the lower Tremadocian (Lower Ordovician) of Liujiachang, south-western Hubei, central China. Palaeontology 58(5), 925–934. View article

Männil, R.M. 1958. New bryozoans of the order Cryptostomata from Ordovician of Estonia. Eesti NSV Teaduste Akadeemia Toimetised, Tehniliste ja füüsikalis-matemaatiliste teaduste seeria 7(4), 330–347. [in Russian] View article

Männil, R.M. 1959. Questions of stratigraphy and bryozoans of the Ordovician of Estonia. 636 pp. Ph.D. thesis, Institut Geologii Akademii Nauk Estonskoi SSR, Tallinn, Estonia. [in Russian]

McKinney, F.K. 1971. Trepostomatous Ectoprocta (Bryozoa) from the lower Chickamauga Group (Middle Ordovician), Wills Valley, Alabama. Bulletins of American Paleontology 60, 195–333.

McKinney, F.K. 1986. Evolution of erect marine bryozoan faunas: repeated success of unilaminate species. The American Naturalist 128, 795–809. View article

McKinney, F.K. & Jackson, J.B.C. 1989. Bryozoan Evolution. Studies in Paleobiology. 238 pp. Unwin Hyman, Boston.

Meidla, T. 1997. Kunda Stage, 64–66. In Raukas, A. & Teedumäe, A. (eds) Geology and Mineral Resources of Estonia. Estonian Academy Publishers, Tallinn.

Modzalevskaya, E.A. 1953. Trepostomata of the Ordovician of East Baltic and their stratgraphic significance. Trudy Vsesojuznogo Nauchnoissledovatelskogo Geologo-Razvedochnogo Instituta (VNIGRI) 78, 91–167. [in Russian]

Modzalevskaya, E.A. 1986. Bryozoa of the family Dittoporidae from the Ordovician of the Leningrad District. Ezhegodnik Vsesojuznogo Paleontologicheskogo Obtchestva 29, 74–91. [in Russian]

Nekhoroshev, V.P. 1961. Ordovician and Silurian bryozoans of Siberian Plate. Trudy VSEGEI, New Series 41, 1–246. [in Russian]

Nelson, C.S., Hyden, F.M., Keane, S.L., Leask, W.L. & Gordon, D.P. 1988. Application of bryozoan zoarial growth-form studies in facies analysis of non-tropical carbonate deposits in New Zealand. Sedimentary Geology 60, 301–322. View article

Nolvak, J., Hints, O. & Männik, P. 2006. Ordovician timescale in Estonia: recent developments. Proceedings of the Estonian Academy of Sciences, Geology 55(2), 95–108. View article

Pushkin, V.I. 1976. New species of Ordovician and Silurian Bryozoa of the Brest Depression, 3–40. In Akimets, V.S. & Govetskii, G.I. (eds) New species of the fossil plants and animals of Belarus. Nauka i Tekhnika, Minsk. [in Russian]

Pushkin, V.I. 1977. About systematic position and the composition of the genus Dittopora (Bryozoa). Transactions of the museum of earth-study of Moscow University 12, 97–113. [in Russian]

Pushkin, V.I. 1987. Bryozoans of suborder Halloporina from Belorussia and East European Platform, 145–232. In Ropot, V.P. & Pushkin, V.I. (eds) Ordovician of Belarus. Nauka i Tekhnika, Minsk. [in Russian]

Pushkin, V.I. 1990. Bryozoa of the family Ceramoporidae of Ordovician from the eastern European platform, 5–34. In New representatives of the fossil fauna and flora of Belorussia and other regions of the USSR: collected scientific papers. Nauka i Tekhnika, Minsk. [in Russian]

Pushkin, V.I. 2002. Middle Ordovician Bryozoa from the Podlasie-Brest Depression (Belarussian part); suborders Ceramoporina, Esthonioporina, Amplexoporina, Halloporina. Geological Quarterly 46(4), 411–434.

Pushkin, V.I. & Popov, L.E. 2001. The Dianulites genus (Bryozoa, Trepostomida) in the Arenig and Llanvirn of the East European Platform. Lithosphere (Litasfera, Belarus) 15, 12–31.

Ross, J.R.P. 1970. Distribution, paleoecology, and correlation of Champlainian Ectoprocta (Bryozoa), New York State, Part III. Journal of Paleontology 44, 346–382.

Schopf, T.J.M., Collier, K.O. & Bach, B.O. 1980. Relation of the morphology of stick-like bryozoans at Friday Harbour, Washington, to bottom currents, suspended matter and depth. Paleobiology 6, 466–476. View article

Silliman, B., Silliman, B., Jr & Dana, J.D. 1851. New genera of fossil corals from the report of James Hall, on the palaeontology of New York. American Journal of Sciences and Arts 2, 398–401.

Smith, A.M., Key, M.M., Jr & Gordon, D.P. 2006. Skeletal mineralogy of bryozoans: taxonomic and temporal patterns. Earth Science Reviews 78(3–4), 287–306. View article

Spjeldnaes, N. 1996. Bryozoan colonies as indicators of bottom conditions in the lower Ordovician, 315–320. In Gordon, D.P., Smith, A.M. & Grant-Mackie, J.A. (eds) Bryozoans in Space and Time. Proceedings of the 10th International Bryozology Conference, Wellington, New Zealand, 1995. National Institute of Water and Atmospheric Research Ltd, Wellington.

Tavener-Smith, R. 1969. Wall structures and acanthopores in the bryozoan Leioclema asperum. Lethaia 2, 89–97. View article

Taylor, P.D. 1975. Monticules in Jurassic cyclostomatous bryozoan. Geological Magazine 112, 601–606. View article

Taylor, P.D. 1979. The inference of extrazooidal feeding currents in fossil bryozoan colonies. Lethaia 12, 47–56. View article

Taylor, P.D. 1999. Bryozoans, 623–645. In Savazzi, E. (ed.) Functional morphology of the invertebrate skeleton. Wiley, New York.

Taylor, P.D. & Wilson, M.A. 1999. Dianulites Eichwald, 1829: an unusual Ordovician bryozoan with a high-magnesium calcite skeleton. Journal of Paleontology 73, 38–48. View article

Torsvik, T. & Cocks, L. 2016. Earth History and Palaeogeography. 332 pp. Cambridge University Press, Cambridge. View article

Ulrich, E.O. 1882. American Palaeozoic Bryozoa. The Journal of the Cincinnati Society of Natural History 5, 121–175, 233–257.

Ulrich, E.O. 1886. Descriptions of new Silurian and Devonian fossils. Bryozoa. Contributions to American Palaeontology 1, 8–33. View article

Ulrich, E.O. 1890. Palaeozoic Bryozoa: III. Report of the Geological Survey of Illinois 8, 283–688.

Ulrich, E.O. 1893. On Lower Silurian Bryozoa of Minnesota. The Geological and Natural History Survey of Minnesota, final report 3, 96–332. View article

Utgaard, J. 1968. A revision of North American genera of ceramoporoid bryozoans (Ectoprocta): Part II; Crepipora, Ceramoporella, Acanthoceramoporella, and Ceramophylla. Journal of Paleontology 42, 1444–1455.

Utgaard, J. 1983. Systematic descriptions for the Order Cystoporata, 357–439. In Robison, R.A. (ed.) Treatise on Invertebrate Paleontology, Part G (1): Bryozoa (revised). Geological Society of Ameri¬ca and Univer¬sity of Kansas Press, Boulder.

Vinassa de Regny, P. 1921. Sulla classificazione dei trepostomidi. Societa Italiana di Sceinze Naturali Atti 59, 212–231.

Vine, G.R. 1884. Fourth report of the Committee appointed for the purpose of reporting on fossil Polyzoa. Reports of the 53rd meeting of the British Association for Advances in Science, 161–209.

Winston, J.E. 1978. Polypide morphology and feeding in marine ectoprocts. Bulletin of Marine Science 28, 1–31.

Winston, J.E. 1979. Current-related morphology and behaviour in some Pacific coast bryozoans, 247–268. In Larwood, G.P. & Abbott, M.B. (eds) Advances in Bryozoology. Academic Press, London.

Xia, F., Zhang, S. & Wang, Z. 2007. The oldest bryozoans: new evidence from the Late Tremadocian (Early Ordovician) of East Yangtze Gorges in China. Journal of Paleontology 81(6), 1308–1326. View article

Yang, J. 1957. Some Bryozoa from the upper part of the Lower Ordovician of Liangshan, southern Shensi (including a new genus). Acta Paleontologica Sinica 5, 1–10.