Abstract
Non-indigenous bryozoans are frequent components of biofouling assemblages in harbour environments worldwide. We performed a survey of artificial hard substrates in six harbours spanning 16 degrees of latitude along the coast of Argentina, from Ingeniero White (38º47′ S) to Ushuaia (54º48′ S). Microeciella argentina n. sp., distributed in warm-temperate waters of northern Patagonia and Buenos Aires Province, is described. The non-indigenous species (NIS) Callopora dumerilii, Smittoidea spinigera and Stephanollona boreopacifica are recorded here for the first time in the Southwest Atlantic. Their occurrence in fouling assemblages of warm-temperate harbours and their previous absence in these areas suggest that they probably arrived in Argentina by international shipping traffic. Callopora dumerilii is native to Europe and the northeast Atlantic. Smittoidea spinigera and Stephanollona boreopacifica are native to China and Korea, respectively. The morphological differences between S. spinigera and the Californian species S. prolifera, which is invading European harbours in the North Sea, are discussed. Bugula neritina, Bugulina flabellata, Cryptosula pallasiana and Fenestrulina delicia, common in harbours of Buenos Aires Province, are here recorded in Patagonia, highlighting the importance of regional shipping traffic to secondary spread of NIS. Arachnidium fibrosum, Buskia socialis, Anguinella palmata, Aetea cultrata, Bicellariella edentata and Exochella moyanoi, previously known from Brazil, and Hippothoa divaricata, are here recorded in Argentina. This study shows that warm-temperate harbours in Buenos Aires Province and northern Patagonia are particularly prone to the introduction of non-indigenous bryozoans.
References
Alder, J. (1856) A catalogue of the zoophytes of Northumberland and Durham. Transactions of the Tyneside Naturalists’ Field Club, 3, 93–160.
Aliotta, S. & Farinati, E. (1990) Stratigraphy of Holocene sand-shell ridges in the Bahia Blanca Estuary, Argentina. Marine Geology, 94, 353–360. https://doi.org/10.1016/0025-3227(90)90064-Q
Allman, G.J. (1856) A monograph of the Freshwater Polyzoa, including all the known species, both British and Foreign. The Ray Society, London, 119 pp. https://doi.org/10.5962/bhl.title.9143
Amor, A. & Pallares, R.E. (1965) Entoprocta y Ectoprocta de la Ría Deseado (Santa Cruz, Argentina) y de otras localidades patagónicas. Physis, Buenos Aires, 25, 291–317.
Audouin, J.V. (1826) Explication sommaire des planches de polypes de l’Egypte et de la Syrie, publiées par Jules-César Savigny. In: Description de l’Egypte, ou receuil des observations et des recherches qui ont été faites en Egypte pendant l’Expedition de l ‘armée française ... Histoire naturelle. Vol. 1. Part 4. Imprimerie Impériale, Paris, pp. 1–339.
Bailey, S.A., Brown, L., Campbell, M.L., Canning-Clode, J., Carlton, J.T., Castro, N., Chainho, P., Chan, F.T., Creed, J., Curd, A., Darling, J., Fofonoff, P., Galil, B.S., Hewitt, C.L., Inglis, G.J., Keith, I., Mandrak, N.E., Marchini, A., McKenzie, C.H., Occhipinti-Ambrogi, A., Ojaveer, H., Pires-Teixeira, L.M., Robinson-Smythe, T., Ruiz, G.M., Seaward, K., Schwindt, E., Son, M., Therriault, T.W. & Zhan, A. (2020) Trends in the detection of aquatic non-indigenous species across global marine, estuarine and freshwater ecosystems: a 50-year perspective. Diversity and Distributions, 26, 1780–1797. https://doi.org/10.1111/ddi.13167
Balavoine, P. (1958) Bryozoaires recueillis a la Nouvelle Amsterdam par M.E. Aubert de la Rue en 1953. Bulletin du Muséum National d’Histoire Naturelle, Série 2, 30, 300–301.
Borg, F. (1926) Studies on Recent cyclostomatous Bryozoa. Zoologiska Bidrag från Uppsala, 10, 181–507.
Bortolus, A. (2008) Error cascades in the biological sciences: The unwanted consequences of using bad taxonomy in ecology. Ambio, 37, 114–118. https://doi.org/10.1579/0044-7447(2008)37[114:ECITBS]2.0.CO;2
Brezina, S.S., Taylor, P.D., Romero, M.V., Palópolo, E.E. & Casadío, S. (2021) Upper Maastrichtian and Danian bryozoans from Northern Patagonia, Argentina. Cretaceous Research, 125, 104845. https://doi.org/10.1016/j.cretres.2021.104845
Busk, G. (1852a) Catalogue of marine Polyzoa in the collection of the British Museum, I. Cheilostomata (part). Trustees of the British Museum, London, 54 pp. https://doi.org/10.1242/jcs.s1-1.2.136b
Busk, G. (1852b) An account of the Polyzoa, and sertularian Zoophytes, collected in the voyage of the Rattlesnake, on the coasts of Australia and the Louisiade Archipelago, &c. In: MacGillivray, J. (Ed.), Narrative of the Voyage of H.M.S. Rattlesnake, commanded by the late Captain Owen Stanley...1846–1850; Including Discoveries and Surveys in New Guinea, the Louisiade Archipelago, etc., to which is added the Account of Mr E.B. Kennedy’s Expedition for the Exploration of the Cape York Peninsula (including Mr W. Carron’s Narrative). T.W. Boone, London, pp. 343–402, pl. 1.
Busk, G. (1859) A monograph of the fossil Polyzoa of the Crag. The Palaeontographical Society, London, 136 pp. https://doi.org/10.5962/bhl.title.2037
Canu, F. (1904) Les bryozoaires du Patagonien. Échelle des bryozoaires pour les terrains tertiaires. Mémoires de la Société Géologique de France, Paléontologie, 12, 1–30.
Canu, F. (1908) Iconographie des bryozoaires fossiles de l’Argentine. Première partie. Anales del Museo Nacional de Buenos Aires, Serie 3, 10, 245–341.
Canu, F. (1911) Iconographie des bryozoaires fossiles de l’Argentine. Deuxième partie. Anales del Museo Nacional de Buenos Aires, Serie 3, 14, 214–288.
Canu, F. (1918) Les ovicelles des bryozoaires cyclostomes. Etudes sur quelques families nouvelles et anciennes. Bulletin de la Société Géologique de France, série 4, 16, 324–335.
Canu, F. & Bassler, R.S. (1925) Les bryozoaires du Maroc et de Mauritanie. Mémoires de la Société des Sciences Naturelles du Maroc, 10, 1–79.
Carlton, J.T. (1996) Biological invasions and cryptogenic species. Ecology, 77, 1653–1655. https://doi.org/10.2307/2265767
Carlton, J.T. (2009) Deep invasion ecology and the assembly of communities in historical time. In: Rilov, G. & Crooks, J.A. (Eds.), Biological invasions in marine ecosystems. Ecological, management, and geographic perspectives. Springer, Berlin and Heidelberg, pp. 13–56. https://doi.org/10.1007/978-3-540-79236-9_2
Carlton, J.T. & Hodder, J. (1995) Biogeography and dispersal of coastal marine organisms: experimental studies on a replica of a 16th-century sailing vessel. Marine Biology, 121, 721–730. https://doi.org/10.1007/BF00349308
Castro, K.L., Giachetti, C.B., Battini, N., Bortolus, A. & Schwindt, E. (2020) Cleaning by beaching: introducing a new alternative for hull biofouling management in Argentina. Aquatic Invasions, 15, 63–80. https://doi.org/10.3391/ai.2020.15.1.05
Castro, K.L., Battini, N., Giachetti, C.B., Trovant, B., Abelando, M., Basso, N.G. & Schwindt, E. (2021) Early detection of marine invasive species following the deployment of an artificial reef: Integrating tools to assist the decision-making process. Journal of Environmental Management, 297, 113333. https://doi.org/10.1016/j.jenvman.2021.113333
Cook, P.L. (1985) Bryozoa from Ghana. A preliminary survey. Annales Musée Royal de l’Afrique Centrale, Sciences Zoologiques, Tervuren, 238, 1–315.
Coutts, S.R., Helmstedt, K.J. & Bennett, J.R. (2018) Invasion lags: The stories we tell ourselves and our inability to infer process from pattern. Diversity and Distributions, 24, 244–251. https://doi.org/10.1111/ddi.12669
Crooks, J.A. & Soulé, M.E. (1999) Lag times in population explosions of invasive species: Causes and implications. In: Sandlund, O.T., Schei, P.J. & Viken, A. (Eds.), Invasive species and biodiversity management. Kluwer Academic Publishers, Dordrecht, pp. 103–125.
De Blauwe, H. (2009) Mosdiertjes van de Zuidelijke bocht van de Noordzee: determinatiewerk voor België en Nederland. Vlaams Instituut voor de Zee, Oostende, 445 pp.
De Blauwe, H. & Faasse, M. (2004) Smittoidea prolifica Osburn, 1952 (Bryozoa, Cheilostomatida), a Pacific bryozoan introduced to The Netherlands (Northeast Atlantic). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Biologie, 74, 33–39.
De Blauwe, H., Kind, B., Kuhlenkamp, R., Cuperus, J., van der Weide, B. & Kerckhof, F. (2014) Recent observations of the introduced Fenestrulina delicia Winston, Hayward and Craig, 2000 (Bryozoa) in Western Europe. In: Rosso, A., Wyse Jackson, P.N. & Porter, J.S. (Eds.), Bryozoan Studies 2013. Studi Trentini di Scienze Naturali, 94, pp. 45–51.
Dick, M.H., Grischenko, A.V. & Mawatari, S.F. (2005) Intertidal Bryozoa (Cheilostomata) of Ketchikan, Alaska. Journal of Natural History, 39, 3687–3784. https://doi.org/10.1080/00222930500415195
Duvergier, J. (1921) Note sur les Bryozoaires du Néogène de l’Aquitaine. Actes de la Société Linnéenne de Bordeaux, 72, 145–182.
Ehrenberg, C.G. (1831) s.n. In: Hemprich, P.C. & Ehrenberg, C.G. (Eds.), Symbolae Physicae, seu Icones et descriptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegyptum, Nubiam, Dongalam, Syriam, Arabiam et Habessiniam...studio annis 1820–25 redirerunt. Ex Officina Academica, venditur a Mittlero, Berolini, pp. 1–162.
Fehlauer-Ale, K.H., Mackie, J.A., Lim-Fong, G.E., Ale, E., Pie, M.R. & Waeschenbach, A. (2013) Cryptic species in the cosmopolitan Bugula neritina complex (Bryozoa, Cheilostomata). Zoologica Scripta, 43, 193–205. https://doi.org/10.1111/zsc.12042
Ferrario, J., d’Hondt, J.-L., Marchini, A. & Occhipinti-Ambrogi, A. (2015) From the Pacific Ocean to the Mediterranean Sea: Watersipora arcuata, a new non-indigenous bryozoan in Europe. Marine Biology Research, 11, 909–919. https://doi.org/10.1080/17451000.2015.1041531
Gabb, W.M. & Horn, G.H. (1862) The fossil Polyzoa of the Secondary and Tertiary formations of North America. Journal of the Academy of Natural Sciences of Philadelphia, 5, 111–179, pls. 19–21.
Geller, J. (1999) Decline of a native mussel masked by sibling species invasion. Conservation Biology, 13, 661–664. https://doi.org/10.1046/j.1523-1739.1999.97470.x
Giachetti, C.B., Battini, N., Castro, K.L. & Schwindt, E. (2020) Invasive ascidians: How predators reduce their dominance in artificial structures in cold temperate areas. Journal of Experimental Marine Biology and Ecology, 533, 151459. https://doi.org/10.1016/j.jembe.2020.151459
Gollasch, S. (2002) The importance of ship hull fouling as a vector of species introductions into the North Sea. Biofouling, 18, 105–121. https://doi.org/10.1080/08927010290011361
Gordon, D.P. (1984) The marine fauna of New Zealand: Bryozoa: Gymnolaemata from the Kermadec Ridge. New Zealand Oceanographic Memoir, 91, 1–198.
Gordon, D.P. (2020) New Hippothoidae (Bryozoa) from Australasia. Zootaxa, 4750 (4), 451–476. https://doi.org/10.11646/zootaxa.4750.4.1
Gordon, D.P. & Mawatari, S.F. (1992) Atlas of marine-fouling Bryozoa of New Zealand ports and harbours. Miscellaneous Publications New Zealand Oceanographic Institute, 107, 1–52.
Gould, A.A. (1850) Description of shells, collected by the North Pacific Exploring Expedition. Proceedings of the Boston Society of Natural History, 3, 151–156 + 169–172 + 214–218 + 252–256 + 275–278 + 292–296 + 309–312 + 343–348.
Gray, J.E. (1848) List of the specimens of British animals in the collection of the British Museum. Part I. Centroniae or radiated animals. Trustees of the British Museum (Natural History), London, 173 pp.
Harmelin, J.-G. (1976) Le sous-ordre des Tubuliporina (Bryozoaires Cyclostomes) en Méditerranée, écologie et systématique. Mémoires de l’Institute Océanographique de Monaco, 10, 1–326.
Harmelin, J.-G. (2014) Alien bryozoans in the eastern Mediterranean Sea—new records from the coast of Lebanon. Zootaxa, 3893 (3), 301–338. https://doi.org/10.11646/zootaxa.3893.3.1
Harmer, S.F. (1915) The Polyzoa of the Siboga Expedition. Part 1. Entoprocta, Ctenostomata and Cyclostomata. Siboga Expedition Reports, 28A, 1–180.
Hastings, A.B. (1927) Zoological results of the Cambridge expedition to the Suez Canal, 1924. Report on the Polyzoa. Transactions of the Zoological Society of London, 22, 331–353. https://doi.org/10.1111/j.1096-3642.1927.tb00381.x
Hastings, A.B. (1943) Polyzoa (Bryozoa) I. Scrupocellariidae, Epistomiidae, Farciminariidae, Bicellariellidae, Aeteidae, Scrupariidae. Discovery Reports, 22, 301–510.
Hayward, P.J. (1985) Ctenostome bryozoans. In: Kermack, D.M. & Barnes, R.S.K. (Eds.), Synopses of the British Fauna. New Series. No. 33. The Linnean Society of London, London, pp. 1–169.
Hayward, P.J. (1991) Systematic studies on some Antarctic and sub-Antarctic Ascophora (Bryozoa: Cheilostomata). Zoological Journal of the Linnean Society, 101, 299–335. https://doi.org/10.1111/j.1096-3642.1991.tb00655.x
Hayward, P.J. & Ryland, J.S. (1985a) Systematic notes on some British Cyclostomata (Bryozoa). Journal of Natural History, 19, 1073–1078. https://doi.org/10.1080/00222938500770671
Hayward, P.J. & Ryland, J.S. (1985b) Cyclostome bryozoans. In: Kermack, D.M. & Barnes, R.S.K. (Eds.), Synopses of the British Fauna. New Series. No. 34. The Linnean Society of London, London, pp. 1–147.
Hayward, P.J. & Ryland, J.S. (1998) Cheilostomatous Bryozoa. Part 1. Aeteoidea—Cribrilinoidea. Synopses of the British Fauna. New Series. No. 10. In: Barnes, R.S.K. & Crothers, J.H. (Eds.), Field Studies Council, Shrewsbury, pp. 1–366.
Hayward, P.J. & Ryland, J.S. (1999) Cheilostomatous Bryozoa. Part 2. Hippothooidea—Celleporoidea. In: Barnes, R.S.K. & Crothers, J.H. (Eds.), Synopses of the British Fauna. New Series. No. 14. Field Studies Council, Shrewsbury, pp. 1–416.
Hincks, T. (1859) On some new and interesting forms of British zoophytes. Quarterly Journal of Microscopical Science, 7, 131–132. https://doi.org/10.1242/jcs.s1-7.26.131a
Hincks, T. (1880) A history of the British Marine Polyzoa. Vol. 1. Van Voorst, London, 601 pp. https://doi.org/10.5962/bhl.title.31555
Hincks, T. (1887) 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, 19, 302–316. https://doi.org/10.1080/00222938709460247
Jullien, J. (1888) Bryozoaires. Mission Scientifique du Cap Horn 1882-1883, 6, 1–92.
Kind, B. & Kuhlenkamp, R. (2018) Discovery of the non-indigenous bryozoan Smittoidea prolifica Osburn, 1952 near Helgoland: first record in 2011 for the German North Sea. Marine Biodiversity, 48, 1237–1240. https://doi.org/10.1007/s12526-016-0544-8
Lamarck, J.-B. (1819) Histoire naturelle des animaux sans vertèbres, 6 (1), 1–343.
Lamouroux, J.V.F. (1812) Extrait d’un mémoire sur la classification des Polypiers coralligènes non entièrement pierreux. Nouveau Bulletin Scientifique de la Société Philosophique, 3, 181–188.
Lamouroux, J.V.F. (1821) Exposition méthodique des genres de l’ordre des polypiers, avec leur description et celles des principales espèces figurées dans 84 planches; les 63 premières apartenant a l’Histoire naturelle des Zoophytes d’Ellis et Solander. V. Agasse, Paris, 115 pp. https://doi.org/10.5962/bhl.title.11328
Levinsen, G.M.R. (1909) Morphological and systematic studies on the cheilostomatous Bryozoa. Nationale Forfatterers Forlag, Copenhagen, 431 pp. https://doi.org/10.5962/bhl.title.5690
Lichtschein de Bastida, V. & Bastida, R. (1980) Los briozoos de las comunidades incrustantes de puertos argentinos. In: Aritio, L. (Ed.), Proceedings of the 5th Intermational Congress on Marine Corrosion and Fouling, Marine Biology, Barcelona, Spain. Gráficas Orbe, Madrid, pp. 371–390.
Lindegren, M., Gabellini, A.P., Munk, P., Edelvang, K. & Hansen, F.T. (2022) Identifying key processes and drivers affecting the presence of non‑indigenous marine species in coastal waters. Biological Invasions, 24, 2835–2850. https://doi.org/10.1007/s10530-022-02802-w
Linnaeus, C. (1758) Systemae naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differetiis, synonymis, locis. 10th Edition. Laurentii Salvii, Holmiae, 795 pp. https://doi.org/10.5962/bhl.title.542
Liu, X. (1990) Three new cheilostome bryozoans from the coasts of Shandong and Zhejiang provinces. Studia Marina Sinica, 31, 121–128.
Liu, X., Yin, X. & Ma, J. (2001) Biology of marine-fouling bryozoans in the coastal waters of China. Science Press, Beijing, 860 pp.
Liuzzi, M.G., López Gappa, J. & Schwindt, E. (2018) Hallazgo de una especie exótica del género Smittoidea (Bryozoa: Cheilostomata) en dos puertos argentinos. In: Libro de Resúmenes X Jornadas Nacionales de Ciencias del Mar. FCEyN-UBA, Buenos Aires, pp. 218.
Lonhart, S.I. (2009) Natural and climate change mediated invasions. In: Rilov, G. & Crooks, J.A. (Eds.), Biological invasions in marine ecosystems. Ecological, management, and geographic perspectives. Springer, Berlin and Heidelberg, pp. 57–70. https://doi.org/10.1007/978-3-540-79236-9_3
López Gappa, J. (1985) Briozoos marinos de la Ría Deseado (Santa Cruz, Argentina). II. Familia Hippothoidae. Physis, Buenos Aires, Secc. A, 43, 51–63.
López Gappa, J. (2000) Species richness of marine Bryozoa in the continental shelf and slope off Argentina (south-west Atlantic). Diversity and Distributions, 6, 15–27. https://doi.org/10.1046/j.1472-4642.2000.00067.x
López-Gappa, J. & Liuzzi, M.G. (2016) High density of the alien bryozoan Fenestrulina delicia in the fouling assemblage of a South American harbour (Argentina). Marine Biodiversity, 46, 509–513. https://doi.org/10.1007/s12526-015-0380-2
López-Gappa, J. & Liuzzi, M.G. (2018) Recent discovery of non-indigenous bryozoans in the fouling assemblage of Quequén Harbour (Argentina, Southwest Atlantic). Marine Biodiversity, 48, 1159–1167. https://doi.org/10.1007/s12526-016-0561-7
Marcus, E. (1937) Bryozoários marinhos brasileiros, 1. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Zoologia, 1, 5–224. https://doi.org/10.11606/issn.2526-4877.bsffclzoologia.1937.113912
Marcus, E. (1938) Bryozoários marinhos brasileiros, 2. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Pauloi, 2, 1–137. https://doi.org/10.11606/issn.2526-4877.bsffclzoologia.1938.113913
Marcus, E. (1939) Bryozoários Marinhos Brasileiros, 3. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Zoologia, 3, 111–353. https://doi.org/10.11606/issn.2526-4877.bsffclzoologia.1939.114101
Marcus, E. (1941) Briozoários marinhos do litoral paranaense. Arquivos do Museu Paranaense, 1, 7–36.
Marcus, E. (1949) Some Bryozoa from the Brazilian coast. Comunicaciones Zoológicas del Museo de Historia Natural de Montevideo, 53, 1–33.
Marcus, E. (1955) Notas sobre briozoos marinhos brasileiros. Arquivos do Museu Nacional do Rio de Janeiro, 42, 273–341.
Markert, A., Matsuyama, K., Rohde, S., Schupp, P. & Wehrmann, A. (2016) First record of the non-native Pacific bryozoan Smittoidea prolifica Osburn, 1952 at the German North Sea coast. Marine Biodiversity, 46, 717–723. https://doi.org/10.1007/s12526-015-0415-8
Maturo, F.J.S. (1968) The distributional pattern of the Bryozoa of the east coast of the United States exclusive of New England. Atti della Societá Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 108, 261–284.
McCann, L.D., McCuller, M.I., Carlton, J.T., Keith, I., Geller, J.B. & Ruiz, G.M. (2019) Bryozoa (Cheilostomata, Ctenostomata, and Cyclostomata) in Galapagos Island fouling communities. Aquatic Invasions, 14, 85–131. https://doi.org/10.3391/ai.2019.14.1.04
McCuller, M.I. & Carlton, J.T. (2018) Transoceanic rafting of Bryozoa (Cyclostomata, Cheilostomata, and Ctenostomata) across the North Pacific Ocean on Japanese tsunami marine debris. Aquatic Invasions, 13, 137–162. https://doi.org/10.3391/ai.2018.13.1.11
McGovern, T.M. & Hellberg, M.E. (2003) Cryptic species, cryptic endosymbionts, and geographical variation in chemical defenses in the bryozoan Bugula neritina. Molecular Ecology, 12, 1207–1215. https://doi.org/10.1046/j.1365-294X.2003.01758.x
Meloni, M., Correa, N., Bettini Pitombo, F., Chiesa, I.L., Doti, B., Elías, R., Genzano, G., Giachetti, C.B., Giménez, D., López-Gappa, J., Pastor, C., Pereira Wandeness, A., Ramírez, F.C., Roccatagliata, D., Schulze-Sylvester, M., Tatián, M., Zelaya, D.G. & Sylvester, F. (2021) In-water and dry-dock hull fouling assessments reveal high risk for regional translocation of nonindigenous species in the southwestern Atlantic. Hydrobiologia, 848, 1981–1996. https://doi.org/10.1007/s10750-020-04345-4
Miranda, A.A., Almeida, A.C.S. & Vieira, L.M. (2018) Non-native marine bryozoans (Bryozoa: Gymnolaemata) in Brazilian waters: Assessment, dispersal and impacts. Marine Pollution Bulletin, 130, 184–191. https://doi.org/10.1016/j.marpolbul.2018.03.023
Molina, J.I. (1782) Saggio sulla storia naturale del Chili. Stamperia di S. Tommaso d’Aquino, Bologna, 367 pp. https://doi.org/10.5962/bhl.title.62689
Moll, J.P.C. von (1803) Eschara, ex zoophytorum, seu, phytozoorum ordine pulcherrimum ac notato dignissimum genus novis speciebus auctum, methodice descriptum et iconibus ad naturam delineatis illustratum. In Libraria Camesiniana, Vindobonae, 70 pp.
Morris, P.A. (1980) The bryozoan family Hippothoidae (Cheilostomata-Ascophora), with emphasis on the genus Hippothoa. Monograph Series of the Allan Hancock Foundation, No. 10, 1–115.
Moyano, H.I. (1982) Magellanic Bryozoa: Some ecological and zoogeographical aspects. Marine Biology, 67, 81–96. https://doi.org/10.1007/BF00397097
Moyano, H.I. (1986) Bryozoa marinos chilenos VI. Cheilostomata Hippothoidae: South Eastern Pacific species. Boletín de la Sociedad de Biología de Concepción, 57, 89–135.
Norman, A.M. (1903) Notes on the natural history of East Finmark. Polyzoa. Annals and Magazine of Natural History, Series 7, 11, 567–598. https://doi.org/10.1080/00222930308678818
Oken, L. (1815) Lehrbuch der Naturgeschichte. Voll. III. Zoologie. Abteilung 1. Fleischlose Thiere, Leipzig, 842 pp.
Orensanz, J.M., Schwindt, E., Pastorino, G., Bortolus, A., Casas, G., Darrigran, G., Elías, R., López Gappa, J.J., Obenat, S., Pascual, M., Penchaszadeh, P., Piriz, M.L., Scarabino, F., Spivak, E.D. & Vallarino, E.A. (2002) No longer the pristine confines of the world ocean: a survey of exotic marine species in the southwestern Atlantic. Biological Invasions, 4, 115–143. https://doi.org/10.1023/A:1020596916153
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. (1933) The Bryozoa of the Mt. Desert region. In: Procter, W. (Ed.), Biological Survey of the Mount Desert region. Wistar Institute of Anatomy and Biology, Philadelphia, Pennsylvania, pp. 3–97.
Osburn, R.C. (1952) Bryozoa of the Pacific coast of America, part 2, Cheilostomata-Ascophora. Report of the Allan Hancock Pacific Expeditions, 14, 271–611.
Penchaszadeh, P.E. (1974) Biología del mejillón Mytilus platensis, su explotación pesquera y perspectivas de cultivo de mitílidos en Argentina. FAO/Carpas, Simposio sobre Acuicultura en América Latina, Montevideo, Uruguay, SE/16, 117–122.
Pereyra, P.J., Narvarte, M., Tatián, M. & González, R. (2015) The simultaneous introduction of the tunicate Styela clava (Herdman, 1881) and the macroalga Undaria pinnatifida (Harvey) Suringar, 1873, in northern Patagonia. BioInvasions Records, 4, 179–184. https://doi.org/10.3391/bir.2015.4.3.04
Pereyra, P.J., de la Barra, P., Gastaldi, M., Saad, J.F., Firstater, F.N. & Narvarte, M.A. (2017) When the tiny help the mighty: facilitation between two introduced species, a solitary ascidian and a macroalga in northern Patagonia, Argentina. Marine Biology, 164, 185. https://doi.org/10.1007/s00227-017-3202-1
Prenant, M. & Bobin, G. (1956) Bryozoaires. Première partie. Entoproctes, Phylactolèmes, Cténostomes. Faune de France, 60, 1–398.
Ramalho, L.V. & Calliari, L. (2015) Bryozoans from Rio Grande do Sul continental shelf, Southern Brazil. Zootaxa, 3955 (4), 569–587. https://doi.org/10.11646/zootaxa.3955.4.8
Reverter-Gil, O., Souto, J. & Fernández-Pulpeiro, E. (2014) Annotated checklist of recent marine Bryozoa from continental Portugal. Nova Acta Científica Compostelana, Bioloxía, 21, 1–55.
Ricciardi, A., Iacarella, J.C., Aldridge, D.C., Blackburn, T.M., Carlton, J.T., Catford, J.A., Dick, J.T.A., Hulme, P.E., Jeschke, J.M., Liebhold, A.M., Lockwood, J.L., MacIsaac, H.J., Meyerson, L.A., Pyšek, P., Richardson, D.M., Ruiz, G.M., Simberloff, D., Vilà, M. & Wardle, D.A. (2021) Four priority areas to advance invasion science in the face of rapid environmental change. Environmental Reviews, 29, 119–141. https://doi.org/10.1139/er-2020-0088
Rilov, G., Benayahu, Y. & Gasith, A. (2004) Prolonged lag in population outbreak of an invasive mussel: a shifting-habitat model. Biological Invasions, 6, 347–364. https://doi.org/10.1023/B:BINV.0000034614.07427.96
Ritter, W.E. & Forsyth, R.H. (1917) Ascidian of the littoral zone of southern California. University of California Publications in Zoology, 16, 439–512.
Rumbold, C., Battini, N., Giachetti, C.B., Castro, K.L., Obenat, S. & Schwindt, E. (2020) Biological invasions in artificial habitats: factors that determine the presence of native and exotic peracarid Crustacea species in Southwestern Atlantic. Marine Biology Research, 16, 340–355. https://doi.org/10.1080/17451000.2020.1781187
Ryland, J.S. (1960) The British species of Bugula (Polyzoa). Proceedings of the Zoological Society of London, 134, 65–105. https://doi.org/10.1111/j.1469-7998.1960.tb05919.x
Ryland, J.S. (1965) Catalogue of main marine fouling organisms, vol 2: Polyzoa. Organisation for Economic Co-operation and Development, Paris, 83 pp.
Ryland, J.S. & Gordon, D.P. (1977) Some New Zealand and British species of Hippothoa (Bryozoa: Cheilostomata). Journal of the Royal Society of New Zealand, 7, 17–49. https://doi.org/10.1080/03036758.1977.10419334
Ryland, J.S., Holt, R., Loxton, J., Spencer Jones, M.E. & Porter, J.S. (2014) First occurrence of the non-native bryozoan Schizoporella japonica Ortmann (1890) in Western Europe. Zootaxa, 3780 (3), 481–502. https://doi.org/10.11646/zootaxa.3780.3.3
Savigny, J.C. (1816) Mémoires sur les animaux sans vertebres. Vol. 2. G. Dufour, Paris, 239 pp. https://doi.org/10.5962/bhl.title.125513
Schwindt, E., Darrigan, G. & Repizo, H. (2010) Evaluación nacional de situación en materia del agua de lastre en el litoral marino y fluvial, Argentina. Proyecto Globallast. Available from: http://archive.iwlearn.net/globallast.imo.org/wp-content/uploads/2015/03/Argentina-Nationalstatus-assessment.pdf (accessed 31 October 2022)
Schwindt, E., López Gappa, J., Raffo, M.P., Tatián, M., Bortolus, A., Orensanz, J.M., Alonso, G., Diez, M.E., Doti, B., Genzano, G., Lagger, C., Lovrich, G., Piriz, M.L., Mendez, M.M., Savoya, V. & Sueiro, M.C. (2014) Marine fouling invasions in ports of Patagonia (Argentina) with implications for legislation and monitoring programs. Marine Environmental Research, 99, 60–68. https://doi.org/10.1016/j.marenvres.2014.06.006
Schwindt, S., Carlton, J.T., Orensanz, J.M., Scarabino, F. & Bortolus, A. (2020) Past and future of the marine bioinvasions along the Southwestern Atlantic. Aquatic Invasions, 15, 11–29. https://doi.org/10.3391/ai.2020.15.1.02
Smitt, F.A. (1868) Kritisk förteckning öfver Skandinaviens Hafs-Bryozoer III. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 24, 279–429.
Taylor, P.D. & Sequeiros, L. (1982) Toarcian bryozoans from Belchite in north-east Spain. Bulletin of the British Museum (Natural History), Geology, 36, 117–129.
Taylor, P.D. & Zaton, M. (2008) Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos (Cyclostomata: Oncousoeciidae). Journal of Natural History, 42, 2557–2574. https://doi.org/10.1080/00222930802277640
van Beneden, P.J. (1845) Récherches sur l’anatomie, la physiologie et le développement des Bryozoaires qui habitent la côte d’Ostende. Nouvelle Memoires de l’Academie Royale des Sciences, des Lettres et des Beaux-arts de Belgique, 18, 1–44.
Viard, F., Roby, C., Turon, X., Bouchemousse, S. & Bishop, J. (2019) Cryptic diversity and database errors challenge non-indigenous species surveys: An illustration with Botrylloides spp. in the English Channel and Mediterranean Sea. Frontiers in Marine Science, 6, 615. https://doi.org/10.3389/fmars.2019.00615
Vieira, L.M., Migotto, A.E. & Winston, J.E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810 (1), 1–39. https://doi.org/10.11646/zootaxa.1810.1.1
Vieira, L.M., Migotto, A.E. & Winston, J.E. (2014) Ctenostomatous Bryozoa from São Paulo, Brazil, with descriptions of twelve new species. Zootaxa, 3889 (4), 485–524. https://doi.org/10.11646/zootaxa.3889.4.2
Vieira, L. M., Almeida, A.C.S. & Winston, J.E. (2016) Taxonomy of intertidal cheilostome Bryozoa of Maceió, northeastern Brazil. Part 1: Suborders Inovicellina, Malacostegina and Thalamoporellina. Zootaxa, 4097 (1), 59–83. https://doi.org/10.11646/zootaxa.4097.1.3
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.
Viviani, C.A. (1977) Bryozoos del litoral chileno. Las especies del género Hippothoa (Ascophora). Medio Ambiente, 2, 38–52.
Waeschenbach, A., Vieira, L.M., Reverter-Gil, O., Souto-Derungs, J., Nascimento, K.B. & Fehlauer-Ale, K.H. (2015) A phylogeny of Vesiculariidae (Bryozoa, Ctenostomata) supports synonymization of three genera and reveals possible cryptic diversity. Zoologica Scripta, 44, 667–683. https://doi.org/10.1111/zsc.12130
Wasson, B. & De Blauwe, H. (2014) Two new records of cheilostome Bryozoa from British waters. Marine Biodiversity Records, 7, e123. https://doi.org/10.1017/S1755267214001213
Wasson, K., Von Holle, B., Toft, J. & Ruiz, G. (2000) Detecting invasions of marine organisms: kamptozoan case histories. Biological Invasions, 2, 59–74. https://doi.org/10.1023/A:1010049907067
Waters, A.W. (1888) Supplementary report on the Polyzoa collected by H.M.S. Challenger during the years 1873–1876. Report on the Scientific Results of the Voyage of the Challenger, Zoology, 31, 1–41.
Waters, A.W. (1904) Bryozoa. Résultats du Voyage du S.V. ‘Belgica’, Zoologie. Expedition Antarctique Belge, 4, 1–114.
Watts, P.C., Thorpe, J.P. & Taylor, P.D. (1998) Natural and anthropogenic dispersal mechanisms in the marine environment: a study using cheilostome Bryozoa. Philosophical Transactions of the Royal Society London B, 353, 453–464. https://doi.org/10.1098/rstb.1998.0222
Whiteaves, J.F. (1901) Catalogue of the marine invertebrata of Eastern Canada. Polyzoa. Geological Survey of Canada, Ottawa, pp. 91–114. https://doi.org/10.4095/216086
Williamson, M. & Fitter, A. (1996) The varying success of invaders. Ecology, 77, 1661–1666. https://doi.org/10.2307/2265769
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. & Hayward, P.J. (2012) The marine bryozoans of the northeast coast of the United States: Maine to Virginia. Virginia Museum of Natural History Memoir, 11, 1–180.
Winston, J.E., Hayward, P.J. & Craig, S.F. (2000) Marine bryozoans of the northeast coast of the United States: New and problem species. In: Herrera-Cubilla, A. & Jackson, J.B.C. (Eds.), Proceedings of the 11th International Bryozoology Association Conference. Smithsonian Tropical Research Institute, Center for Tropical Paleoecology and Archeology, Balboa, pp. 412–420.
Xavier, E.A., Almeida, A.C.S. & Vieira, L.M. (2021) The role of artificial habitats on fouling bryozoan fauna in the southwestern Atlantic. Marine Pollution Bulletin, 167, 112310. https://doi.org/10.1016/j.marpolbul.2021.112310
Yang, H.J., Seo, J.E. & Gordon, D.P. (2018) Sixteen new generic records of Korean Bryozoa from southern coastal waters and Jeju Island, East China Sea: evidence of tropical affinities. Zootaxa, 4422 (4), 493–518. https://doi.org/10.11646/zootaxa.4422.4.3