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The identity of the invasive fouling bryozoan Watersipora subtorquata (d’Orbigny) and some other congeneric species

Leandro M. Vieira, Mary Spencer Jones, Paul D. Taylor


Watersipora subtorquata (d’Orbigny, 1852) has been widely reported as a fouling species from tropical to temperate waters. The continued confusion over the correct name for this species led us to provide a redescription of d’Orbigny’s type of Cellepora subtorquata, and to make comparisons with other species of Watersipora. We show that the majority of specimens assigned to W. subovoidea (d’Orbigny, 1852) are morphologically distinct from the recently erected neotype of W. subovoidea; these specimens are here reidentified as Watersipora subtorquata. Other specimens previously assigned to W. subtorquata belong to W. subatra (Ortmann, 1890), described originally from Japan. Owing these inconsistences, we suggest setting aside the neotype of Watersipora subovoidea, which is based on Busk’s Lepralia cucullata and is not from the same locality as d’Orbigny’s type. Watersipora cucullata is redescribed and figured using Busk’s specimens; the species is known from the Mediterranean, including the Adriatic. Three other species—Watersipora atrofusca (Busk, 1856), Watersipora aterrima (Ortmann, 1890) and Watersipora nigra (Canu & Bassler, 1930)—are also refigured. Watersipora edmondsoni Soule & Soule, 1975 is synonymised with W. subtorquata (d’Orbigny). Two new species are described, Watersipora mawatarii n. sp. from Japan and Watersipora souleorum n. sp. from the Azores, Cape Verde, Naples and Indian Ocean. A key is given to the Recent species of Watersipora.



Bryozoa, Cheilostomata, Watersiporidae, fouling species, invasive species, new species, taxonomy, type specimens

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Abdel-Salam, K.M. & Ramadan, S.E. (2008) Fouling Bryozoa from some Alexandria harbours, Egypt. (II) Encrusting species. The Mediterranean Marine Science Journal, 9 (2), 5–20.

Audouin, J.V. (1826) Explication sommaire des planches de Polypes de l’Egypte et de la Syrie, publiées par Jules-César Savigny. In: Panckoucke, C.L.F. (Org.), Description de l’Egypte ou receuil des observations et des recherches qui ont été faites en Egyptes pendant l’Expedition de l’Armée française … Histoire naturelle. Tome 1(4). Imprimerie Impériale, Paris, pp. 225–244.

Banta, W.C. (1969a) Watersipora arcuata, a new species in the subovoidea-cucullata-nigra complex (Bryozoa, Cheilostomata). Bulletin of the Southern California Academy of Science, 68, 96–102.

Banta, W.C. (1969b) Uscia mexicana, new genus, new species, a watersiporid bryozoan with dimorphic autozooids. Bulletin of the Southern California Academy of Science, 68, 20–35.

Banta, W.C. (1970) The body wall of cheilostome Bryozoa, III. The frontal wall of Watersipora arcuata Banta, with a revision of the Cryptocystidea. Journal of Morphology, 131, 37–56.

Bock, P. (2010) Watersipora Neviani, 1896. In: Recent and Fossil Bryozoa. Mount Waverley (Vic): Philip Bock. Available from: (accessed 20 June 2014)

Busk, G. (1854) Catalogue of marine Polyzoa in the collection of the British Museum, II. Cheilostomata (part). Trustees of the British Museum (Natural History), London, 54 pp.

Busk, G. (1856) Zoophytology. Quarterly Journal of Microscopical Science, 4, 176–179, pls 7–8.

Canu, F. & Bassler, R.S. (1923) North American later Tertiary and Quaternary Bryozoa. United States National Museum Bulletin, 125, 1–302.

Canu, F. & Bassler, R.S. (1930) The Bryozoa of the Galapagos Islands. Proceedings of the United States National Museum, 76, 1–78.

Cockrell, M.L. & Sorte, C.J.B. (2013) Predicting climate-induced changes in population dynamics of invasive species in a marine epibenthic community. Journal of Experimental Marine Biology and Ecology, 440, 42–48.

Cook, P.L. (1985) Bryozoa from Ghana. Tervuren, België Koninklijk Museum voor Midden-Afrika. Zoologische Weteschappen, 238, 1–315.

Davis, K. & Marshall, D.J. (2014) Offspring size in a resident species affects community assembly. Journal of Animal Ecology, 83, 322–331.

Dick, M.H., Tilbrook, J. & Mawatari, S.F. (2007) Diversity and taxonomy of rocky-intertidal Bryozoa on the Island of Hawaii, USA. Journal of Natural History, 40, 2197–2258.

Florence, W.K., Hayward, P.J. & Gibbons, M.J. (2007) Taxonomy of shallow-water Bryozoa from the west coast of South Africa. African Natural History, 3, 1–58.

Geller, J., Mackie, J.A., Schroeder, G. & Gerhinger, D. (2008) Distribution of highly invasive bryozoans belonging to a cryptic species complex in the genus Watersipora determined by DNA sequences. Moss Landing Marine Laboratories, Moss Landing, 40 pp.

Gordon, D.P. (1989) The marine fauna of New Zealand: Bryozoa: Gymnolaemata (Cheilostomida Ascophorina) from the western South Island continental shelf and slope. New Zealand Oceanographic Institute Memoir, 97, 1–158.

Gordon, D.P. & Mawatari, S.H. (1992) Atlas of marine-fouling Bryozoa of New Zealand ports and harbours. Miscellaneous Publications of the New Zealand Oceanographic Institute, 107, 1–52.

Harmer, S.F. (1957) The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga-Expeditie, 28d, xv, 641–1147, pls. 42–74.

Hastings A.B. (1927) Zoological results of the Cambridge expedition to the Suez Canal, 1924, 20. Report on the Polyzoa. Transactions of the Zoological Society of London, 22, 331–353.

Hastings, A.B. (1930) Cheilostomatous Polyzoa from the vicinity of the Panama Canal collected by Dr. A. Crossland on the cruise of the S.Y. 'St. George'. Proceedings of the Zoological Society of London, 99, 697–740. [1929 on cover]

Hayward, P.J. & McKinney, F.K. (2002) Northern Adriatic Bryozoa from the vicinity of Rovinj, Croatia. Bulletin of the American Museum of Natural History, 270, 1–139.<0001:NABFTV>2.0.CO;2

Hincks, T. (1886) The Polyzoa of the Adriatic: a supplement to Prof. Heller’s ‘Die Bryozoen des adriatischen Meeres’, 1867. Annals and Magazine of Natural History, Series 5, 17, 254–271.

Hondt, J. L. d’ (1988) Bryozoa from the coast of Israel. Italian Journal of Zoology, 55, 191–203.

ICZN (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, xx + 365 pp.

Jullien, J. & Calvet, L. (1903) Bryozoaires provenant des campagnes de l’Hirondelle (1886–1888). Résultats des campagnes scientifiques accomplies sur son yacht par Albert 1er, Prince Souverain de Monaco, 1903, 23, 1–188.

Kelso, A. & Wyse Jackson, P.N. (2012) Invasive bryozoans in Ireland: first record of Watersipora subtorquata (d’Orbigny, 1852) and and extension of the range of Tricellaria inopinata d’Hondt and Occhipinti Ambrogi, 1985. BioInvasions Records, 1, 209–214.

Kuhlenkamp, R. & Kind, B. (2013) Arrival of the invasive Watersipora subtorquata (Bryozoa) at Helgoland (Germany, North Sea) on floating macroalgae (Himanthalia). Marine Biodiversity Records, 6, e73. S1755267213000481

Kubota, K. & Mawatari, S.F. (1985) A systematic study of cheilostomatous bryozoans from Oshoro Bay, Hokkaido. 2. Ascophora. Environmental Science, Hokkaido University, 8 (2), 195–208.

Lamouroux, J.V.F. (1816) Histoire des Polypiers coralligènes flexibles, vulgairement nommés Zoophytes. F. Poisson, Caen, 559 pp.

Lamouroux, J.V.F. (1824–26) Description des polypiers flexibles. In: Quoy, J.R.C. & Gaimard, J.P. (Eds.), Voyage autour du Monde … exécuté sur l’Uranie et la Physicienne pendant les années 1817–1820. Vol. 3. Zoologie IV. Pilet Ainé, Paris, pp. 603–642.

Láruson, A.J., Craig, S.F., Messer, K.J. & Mackie, J.A. (2012) Rapid and reliable inference of mitochondrial phylogroups among Watersipora species, and invasive group of ship-fouling species (Bryozoa, Cheilostomata). Conservation Genetics Resources, 4, 617–619.

Mackie, J.A., Keough, M.J. & Christidis, L. (2006) Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata, and Watersipora arcuata. Marine Biology, 149, 285–295.

Mackie, J.A., Darling, J.A. & Geller, J.B. (2012) Ecology of cryptic invasions: latitudinal segregation among Watersipora (Bryozoa) species. Scientific Reports, 2 (871), 1–10.

Manzoni, A. (1875) I briozoi del pliocene antico di Castrocaro. Gamberini & Parmeggiani, Bologna, 64 pp.

Marcus, E. (1937) Bryozoários Marinhos Brasileiros, 1. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Zoologia, 1, 5–224.

Marcus, E. (1938) Bryozoários Marinhos Brasileiros, 2. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Zoologia, 2, 1–196.

Marcus, E. (1955) Notas Sobre Briozoos Marinhos Brasileiros. Arquivos do Museu Nacional do Rio de Janeiro, 42, 273–341.

Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14–17.

Needles, L.A. & Wendt, D.E. (2013) Big changes to a small bay: introduced species and long-term compositional shifts to the fouling community of Morro Bay (CA). Biological Invasions, 15, 1231–1251.

Neviani, A. (1896) Briozoi fossili della Farnesina e Monte Mario presso Roma. Palaeontographia italica, 1, 77–140.

Norman, A.M. (1864) On undescribed British Hydrozoa, Actinozoa and Polyzoa. Annals and Magazine of Natural History, Series 3, 13, 82–90.

Okada, Y. & Mawatari, S. (1937) On the collection of Bryozoa along the coast of Onagawa bay and its vicinity, the northern part of Honshu, Japan. Science Reports of the Tôhoku Imperial University, Biology, 11, 433–445.

Orbigny, A. d’ (1841–1847) Zoophytes. Voyage dans L`Amerique Méridionale, 5(4), 7–28.

Orbigny, A. d’ (1851–1854) Paléontologie française. Description des Mollusques et Rayonnés fossiles. Terrains crétacés, V. Bryozoaires. Victor Masson, Paris, pp. 1–188 (1851); pp. 185 bis–472 (1852); pp. 473–984 (1853); pp. 985–1192 (1854); pls 600–800 (1899). [Dates as given by Sherborn]

Ortmann, A. (1890) Die Japanische Bryozoenfauna. Bericht über die von Herrn Dr. L. Döderlein im Jahre 1880–81 gemachten Sammlungen. Archiv für Naturgeschichte, 54, 1–74.

Osburn, R.C. (1952) Bryozoa of the Pacific coast of America, part 2, Cheilostomata–Ascophora. Allan Hancock Pacific Expeditions, 14, 271–611.

Quoy, J.R.C. & Gaimard, J.P. (1827) Observations zoologiques faites à bord de l’Astrolabe, en mai 1826, dans le détroit de Gibraltar. Annales des Sciences Natureles, Series 1, 10, 1–239.

Ramalho, L.V., Muricy, G. & Taylor, P.D. (2011) Taxonomic revision of some lepraliomorph cheilostome bryozoans (Bryozoa: Lepraliomorpha) from Rio de Janeiro State, Brazil. Journal of Natural History, 45, 757–798.

Ryland, J.S. (1974) Bryozoa in the Great Barrier Reef province. In: Cameron, A.M. et al. (Eds.), Proceedings of the Second International Reef Symposium. Vol. 2. The Great Barrier Reef Committee, Brisbane, pp. 341–348.

Ryland, J.S., de Blauwe, H., Lord, R. & Mackie, J.A.A. (2009) Recent discoveries of alien Watersipora (Bryozoa) in Western Europe, with redescriptions of species. Zootaxa, 2093, 43–59.

Savigny, J.C. (1817) Description de l’Égypte, ou receuil des observations et des recherches qui ont été faites en Égypte pendant l’Expedition de ‘Armée française ... Histoire naturelle. Vol. 2. Imprimerie Royale, Paris, 125 pls.

Seo, J.E. (1999) Taxonomic review of Korean Watersipora (Bryozoa, Gymnolaemata, Cheilostomata). Korean Journal of Systematic Zoology, 15, 221–229.

Seo, J.E. (2005) Illustrated Encyclopedia of Fauna and Flora of Korea. Vol. 40. Bryozoa. Ministry of Education and Human Resources Development, Daehan Printing and Publishing, Seoul, 596 pp.

Sherborn, C.D. (1899) On the dates of the “Paléontologie Française” of d’Orbigny. Geological Magazine, Series 4, 6, 223–225.

Sorte, C.J.B. & White, J.W. (2013) Competitive and demographic leverage points of community shits under climate warming. Proceedings of the Royal Society B, 280, 20130572.

Soule, D.F. & Soule, J.D. (1968) Bryozoan fouling organisms from Oahu, Hawaii, with a new species of Watersipora. Bulletin of the Southern California Academy of Science, 67, 203–218.

Soule, D.F. & Soule, J.D. (1975) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Géologie de la Faculté des Science de Lyon, h.s. 3, 2, 299–309.

Soule, J.D. (1961) Results of the Puritan-American Museum of Natural History Expedition to Western Mexico 13. Ascophoran Cheilostomata (Bryozoa) of the Gulf of California. American Museum Novitates, 2053, 1–66.

Stefani, C. de (1884) Escursione scientifica nella Calabria (1877–78): Jejo, Montalto e Capo Vaticano. Memorie dela Regia Accademia di Scienze Fisiche, Matematiche e Naturali, Series 3, 18, 1–292.

Taylor, P.D. & Gordon, D.P. (2002) Alcide d’Orbigny’s work on Recent and fossil bryozoans. Compte Rendu Palévol, 1, 533–547.

Thornely, L.R. (1905) Report on the Polyzoa collected by Professor Herdmann, at Ceylon, in 1902. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, 4, 107–130.

Tilbrook, K.J., Hayward, P.J. & Gordon, D.P. (2001) Cheilostomatous Bryozoa from Vanuatu. Zoological Journal of the Linnean Society, 131, 35–109.

Tilbrook, K.J. (2006) Cheilostomatous Bryozoa from the Solomon Islands. Santa Barba Museum of Natural History Monographs, 4 (Studies in Biodiversity Number 3), 1–386.

Vieira, L.M., Migotto, A.E. & Winston, J.E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1–39.

Vigneaux, M. (1949) Révision des Bryozoaires néogènes du Bassin d’Aquitaine et essai de classification. Mémoires de la Société Géologique de France, New Series, 28, 1–153.

Waters, A.W. (1879) III.—On the Bryozoa (Polyzoa) of the Bay of Naples. Annals and Magazine of Natural History, ser. 5, 3, 28–43.

Waters, A.W. (1905) Notes on some Recent Bryozoa in d’Orbigny’s collection. Annals and Magazine of Natural History, Series 7, 15, 1–16.

Waters, A.W. (1909) Reports on the Marine Biology of the Sudanese Red Sea, from collections made by Cyril Crossland, M.A., B.Sc., F.Z.S.; together with collections made in the Red Sea by Dr. Hartmeyer. XII. The Bryozoa, Part 1, Cheilostomata. Linnean Society of London Journal of Zoology, 31, 123–181.

Winston, J.E. (1982) Marine bryozoans (Ectoprocta) of the Indian River area, Florida. Bulletin of the American Museum of Natural History, 173, 99–176.

Winston, J.E. & Heimberg, B.F. (1986) Bryozoans from Bali, Lombok, and Komodo. American Museum Novitates, 2847, 1–49.


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