Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2020-03-03
Page range: 201–252
Abstract views: 143
PDF downloaded: 4

Taxonomy and diversity of coelobite bryozoans from drift coral cobbles on
Co To Island, northern Vietnam

Department of Biological Sciences, Faculty of Science, Hokkaido University, N10 W8, Sapporo 060-0810, Japan.
Institute of Marine Environment and Resources, Vietnam Academy of Science and Technology, Hai Phong City, Vietnam.
Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Ha Noi, Vietnam.
Bryozoa competition coral reef encrusting Foraminifera marine Bryozoa new species zoogeography

Abstract

There has been no previous report detailing the taxonomy of marine bryozoans along the coasts of Vietnam. Here we report on the taxonomy and diversity of bryozoans collected among drift coral cobbles from a beach in the tropical Co To archipelago, Gulf of Tonkin, northern Vietnam. We detected 27 bryozoan species (23 cheilostomes, four cyclostomes) in a coelobite assemblage inhabiting crevices in the cobbles, and holes made by boring molluscs. The degree of bryozoan preservation varied greatly, suggesting that the cobbles had accumulated on the beach over a period of months to years, or even decades. Coral reefs in the Co To archipelago underwent a catastrophic decline in 2003–2008, and it is unclear whether the bryozoan assemblage reflects past diversity, present diversity remaining in the coral rubble, or both. We describe six new species: Parasmittina acondylata n. sp., Metroperiella cotoensis n. sp., Microporella tonkinensis n. sp., Rhynchozoon setiavicularium n. sp., R. latiavicularium n. sp., and Disporella phaohoa n. sp. All but two of the previously described species were already known from the Central Indo-Pacific coastal biogeographical realm of Spalding et al. (2007), which includes Vietnam. We report the third Recent record of the thalamoporellid Dibunostoma reversum (Harmer, 1926), which is quite similar to and might be conspecific with the lower Miocene species Thalamoporella transversa Guha & Krishna, 2004; while it is premature to synonymize the two, we transfer T. transversa to Dibunostoma, as D. transversum. The calcareous, sheet-like, encrusting foraminiferan Planorbulina larvata was prominent in the coelobite assemblage and was often observed in substrate competition with bryozoans. A limited analysis of competitive interactions indicated that the encrusting bryozoans in the coelobite assemblage encountered P. larvata more often than they encountered other bryozoans, and that P. larvata out-competed bryozoans for substrate, reinforcing a growing sense of the importance of encrusting foraminifera in tropical and subtropical hard-substrate communities.

 

References

  1. Allmann, G.J. (1856) A Monograph of the Freshwater Polyzoa, Including All the Known Species, both British and Foreign. The Ray Society, London, 119 pp., 11 pl.

    https://doi.org/10.5962/bhl.title.9143

    Audouin, J.V. (1826) Explication sommaire des planches de polypes de l’Égypte et de la Syrie, publiées par Jules-César Savigny [Planches 1–14 by J-C Savigny (1817)]. In: Audouin J.V. (Ed.), Description de l’Égypte, ou Receuil des Observations et des Recherches qui ont été Faites en Égypte Pendant l’Expedition de l’Armée Française, Publié par les Orders de sa Majesté l’Empéreur Napoléon le Grand. Histoire naturelle. Tome 1. 4e Partie. Imprimérie Impériale, Paris, pp. 225–244.

    Badve, R.M. & Sonar, M.A. (1997) Some fossil neocheilostomine bryozoans from the Holocene of the west coast of Maharashtra and Goa, India. Journal of the Palaeontological Society of India, 42, 3548.

    Balavoine, P. (1959) Bryozoaires. Mission Robert Ph. Dollfus en Égypte (décembre 1927 – mars 1929). S.S. “Al Sayad”. Résultats scientifiques, 3e Partie, 34, 257282.

    Berning, B. (2012) Taxonomic notes on some Cheilostomata (Bryozoa) from Madeira. Zootaxa, 3236, 3654.

    https://doi.org/10.11646/zootaxa.3236.1.2

    Bock, P. (2018) Indexes to bryozoan taxa. Available from: http://www.bryozoa.net/indexes.html (accessed 15 December 2018)

    Borg, F. (1926) Studies on Recent cyclostomatous Bryozoa. Zoologiska Bidrag från Uppsala, 10, 181507.

    Brady, H.B. (1884) Report on the Foraminifera dredged by H. M. S. Challenger during the years 1873–1876. Report on the Scientific Results of the Voyage of the H.M.S. Challenger, Zoology, 9, 1814, pls. 1115.

    Bronn, H.G. (1825) System der Urweltlichen Pflanzenthiere Durch Diagnose, Analyse und Abbildung der Geschlechter Erlautert. J.C.B. Mohr, Heidelberg, 47 pp.

    https://doi.org/10.5962/bhl.title.9094

    Busk, G. (1852) 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]. Vol. 1. T. & W. Boone, London, pp. 343–402, pl. 1.

    Busk, G. (1859) A Monograph of the Fossil Polyzoa of the Crag. The Palaeontological Society, London, 136 pp.

    https://doi.org/10.5962/bhl.title.2037

    Buss, L.W. (1979) Bryozoan overgrowth interactions—the interdependence of competition for space and food. Nature, 281, 475–477.

    https://doi.org/10.1038/281475a0

    Buss, L.W. (1980) Competitive intransitivity and size-frequency distributions of interacting populations. Proceedings of the National Academy of Sciences of the United States of America, 77, 535–5359.

    https://doi.org/10.1073/pnas.77.9.5355

    Buss, L.W. (1990) Competition within and between encrusting clonal invertebrates. Trends in Ecology and Evolution, 5, 352–356.

    https://doi.org/10.1016/0169-5347(90)90093-S

    Cáceres-Chamizo, J.P., Sanner, J., Tilbrook, K.J. & Ostrovsky, A.N. (2017) Revision of the Recent species of Exechonella Canu and Bassler in Duvergier, 1924 and Actisecos Canu & Bassler, 1927 (Bryozoa, Cheilostomata): systematics, biogeography and evolutionary trends in skeletal morphology. Zootaxa, 4305 (1), 1–79.

    https://doi.org/10.11646/zootaxa.4305.1.1

    Canu, F. (1918) Les ovicelles des Bryozoaires cyclostomes. Étude sur quelques familles nouvelles et anciennes. Bulletin de la Societé géologique de France, 16 (4), 324­–335.

    Canu, F. & Bassler, R.S. (1917) A synopsis of American Early Tertiary cheilostome Bryozoa. United States National Museum Bulletin, 96, 1–87.

    https://doi.org/10.5479/si.03629236.96.1

    Canu, F. & Bassler, R.S. (1927) Bryozoaires des Isles Hawaï. Bulletin de la Société des Sciences de Seine-et-Oise, 7 (Supplement), 1–67.

    Canu, F. & Bassler, R.S. (1929) Bryozoa of the Philippine region. United States National Museum Bulletin, 100, I–XI + 1–685.

    Canu, F. & Bassler, R.S. (1933) The bryozoan fauna of the Vincentown Limesand. United States National Museum Bulletin, 165, 1108.

    https://doi.org/10.5479/si.03629236.165.i

    Cheetham, A.H. (1963) Late Eocene zoogeography of the eastern Gulf Coast region. Memoirs of the Geological Society of America, 91, 1113.

    https://doi.org/10.1130/MEM91-p1

    Chimonides, P.J. & Cook, P.L. (1994) Notes on the genus Cranosina (Bryozoa, Cheilostomida). Zoologica Scripta, 23, 4349.

    https://doi.org/10.1111/j.1463-6409.1994.tb00372.x

    Defrance, J.L.M. (1823) Polypiers. In: Dictionnaire des Sciences Naturales. Vol. 26. F.G. Levrault, Strasbourg, pp. 1–553.

    Dick, M.H. & Grischenko, A.V. (2017) Rocky-intertidal cheilostome bryozoans from the vicinity of the Sesoko Biological Station, west-central Okinawa, Japan. Journal of Natural History, 51, 141–266.

    https://doi.org/10.1080/00222933.2016.1253797

    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

    Dick, M.H. & Mawatari, S.F. (2005) Morphological and molecular concordance of Rhynchozoon clades (Bryozoa, Cheilostomata) from Alaska. Invertebrate Biology, 124, 344–354.

    https://doi.org/10.1111/j.1744-7410.2005.00032.x

    Dick, M.H. & Ross, J.R.P. (1988) Intertidal Bryozoa (Cheilostomata) of the Kodiak Vicinity, Alaska. Center for Pacific Northwest Studies Occasional Paper 23. Western Washington University, Bellingham, 133 pp.

    Dick, M.H., Tilbrook, K.J. & Mawatari, S.F. (2006) Diversity and taxonomy of rocky-intertidal Bryozoa on the Island of Hawaii, USA. Journal of Natural History, 40, 2197–2257.

    https://doi.org/10.1080/00222930601062771

    Duvergier, J. (1924) Deuxième note sur les Bryozoaires du Néogène de l’Aquitaine. Actes de la Société linnéenne Bordeaux, 75, 145–190.

    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.

    Gluhak, T., Lewis, J. & Popijac, A. (2007) Bryozoan fauna of Green Island, Taiwan: first indications of biodiversity. Zoological Studies, 46, 397–426.

    Gordon, D.P. (2016) Bryozoa of the South China Sea—an overview. Raffles Bulletin of Zoology, Supplement, 34, 604–618.

    Gordon, D.P. (1984) The marine fauna of New Zealand: Bryozoa: Gymnolaemata from the Kermadec ridge. New Zealand Oceanographic Institute Memoir, 91, 1–198.

    Gordon, D.P. (2017) Bryozoa: Cheilostomata. Interim classification for Treatise. Version 8 September 2017. Unpublished but widely circulated document, obtained from D.P. Gordon, National Institute of Water & Atmospheric Research, Wellington, 16 pp.

    Gordon, D.P., Hossain, Md.M.M. & Wood, T.S. (2007) The known and anticipated bryozoan diversity of Bangladesh. Journal of Taxonomy and Biodiversity Research, 1, 45–58.

    Gordon, D.P. & Taylor, P.D. (1997) The Cretaceous-Miocene genus Lichenopora (Bryozoa), with a description of a new species from New Zealand. Bulletin of the Natural History Museum, London, Geology, 53, 71–78.

    Gordon, D.P. & Taylor, P.D. (2001) New Zealand Recent Densiporidae and Lichenoporidae (Bryozoa: Cyclostomata). Species Diversity, 6, 243–290.

    https://doi.org/10.12782/specdiv.6.243

    Gray, J.E. (1848) List of the Specimens of British Animals in the Collections of the British Museum. Part 1. Centrionae or Radiated Animals. Trustees of the British Museum, London, pp. 91–151. [Polyzoa]

    Guha, A.K. & Krishna, K.G. (2004) Fossil Thalamoporella (Bryozoa) from the Tertiary Sequences of Western Kachchh, Gujarat, India. Irene McCulloch Foundation Monograph Series, 7, 1–51.

    Harmelin, J.-G. (1979) On some stomatoporiform species (Bryozoa Cyclostomata) from the bathyl zone of the northeastern Atlantic Ocean. In: Larwood, G.P. & Abbott, M.B, (Eds.), Advances in Bryozoology. Academic Press, London, pp. 403–422.

    Harmelin, J.-G., Ostrovsky, A.N., Cáceres-Chamizo, J.P. & Sanner, J. (2011) Bryodiversity in the tropics: taxonomy of Microporella species (Bryozoa, Cheilostomata) with personate maternal zooids from Indian Ocean, Red Sea and southeast Mediterranean. Zootaxa, 2798 (1), 1–30.

    https://doi.org/10.11646/zootaxa.2798.1.1

    Harmelin, J.-G., Vieira, L.M., Ostrovsky, A.N., Cáceres-Chamizo, J.P. & Sanner, J. (2012) Scorpiodinipora costulata (Canu & Bassler, 1929) (Bryozoa, Cheilostomata), a taxonomic and biogeographic dilemma: complex of cryptic species or human-mediated cosmopolitan colonizer? Zoosystema, 34, 123–138.

    https://doi.org/10.5252/z2012n1a5

    Harmer, S.F. (1926) The Polyzoa of the Siboga Expedition, part 2, Cheilostomata Anasca. Siboga Expedition Reports, 28b, v–vii + 183–501, pls. 13–34.

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

    Hayward, P.J. (1988) Mauritian cheilostome Bryozoa. Journal of the Zoological Society of London, 215, 269–356.

    https://doi.org/10.1111/j.1469-7998.1988.tb04900.x

    Hincks, T. (1877) On British Polyzoa—Part II. Classification. Annals and Magazine of Natural History, Series 4, 20, 520–532.

    https://doi.org/10.1080/00222937708682275

    Hincks, T. (1879) On the classification of the British Polyzoa. Annals and Magazine of Natural History, Series 5, 3, 153–164.

    https://doi.org/10.1080/00222937908682494

    Hincks, T. (1880) Contributions towards a general history of the marine Polyzoa. II. Foreign Membraniporina. Annals and Magazine of Natural History, Series 5, 6, 81–92, pls. 9–11.

    https://doi.org/10.1080/00222938009458895

    Hincks, T. (1881) Contributions towards a general history of the marine Polyzoa. IV. Foreign Membraniporina (second series). Annals and Magazine of Natural History, Series 5, 7, 147–156, pls. 8–10.

    https://doi.org/10.1080/00222938109459489

    Hincks, T. (1882) Contributions towards a general history of the marine Polyzoa. IX. Foreign Cheilostomata (Miscellaneous). Annals and Magazine of Natural History, Series 5, 9, 116–127, pl. 5.

    https://doi.org/10.1080/00222938209459003

    Hincks, T. (1895) Index [to “Marine Polyzoa: Contributions Towards a General History”]. Issued Privately, London, 6 pp.

    Illies, G. (1973) Different budding patterns in the genus Stomatopora (Bryozoa, Cyclostomata). In: Larwood, G.P. (Ed.), Living and Fossil Bryozoa. Academic Press, London, pp. 307–315.

    Johnston, G. (1838) A History of British Zoophytes. W.H. Lizars, Edinburgh, 341 pp.

    https://doi.org/10.5962/bhl.title.110844

    Jullien, J. (1882) Note sur une nouvelle division des Bryozoaires Cheilostomiens. Bulletin de la Société zoologique de France, 6, 271–285.

    Jullien, J. (1888) Bryozoaires. Mission Scientifique du Cap Horn 1882-1883, Series 6, 3, 1–92, 15 pls.

    Kubanin, A.A. (1990) Biogeographical and morphological peculiarities of Bryozoa of the coastal waters of Vietnam. 8th All-Russian Colloquium on Fossil and Modern Bryozoa, Tallinn, Estonia, 1990, 27–29. [in Russian]

    Lamarck, J.B.P.A. de M de. (1816) Histoire naturelle des Animaux sans Vertèbres. Vol. 2. Verdiere, Paris, 568 pp.

    Lamouroux, J.V.F. (1821) Exposition Méthodique des Genres de l’Ordre des Polypiers, avec leur Description et Celles des Principales Espèces, Figureés dans 84 Planches; les 63 Premières Apartenant à 1ʹHistoire Naturelle des Zoophytes dEllis et Solander. V. Agasse, Paris, 115 pp., 84 pls.

    https://doi.org/10.5962/bhl.title.11328

    Levinsen, G.M.R. (1902) Studies on Bryozoa. Videnskabelige Meddelelser fra den naturhistoriske Foreningi København, 54, 1–31.

    Levinsen, G.M.R. (1909) Morphological and Systematic Studies on the Cheilostomatous Bryozoa. Nationale Forfatteres Forlag, Copenhagen, 431 pp.

    https://doi.org/10.5962/bhl.title.5690

    Linnaeus, C. (1758) Systemae Naturae per Regna tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differetiis, Synonymis, Locis. 10th Edition. Laurentii Salvii, Holmiae, iii + 824 pp.

    https://doi.org/10.5962/bhl.title.542

    Liow, L.H., Di Martino, E., Krzeminska, M., Ramsfjell, M., Rust, S., Taylor, P.D. & Voje, K.L. (2017) Relative size predicts competitive outcome through 2 million years. Ecology Letters, 20, 981–988.

    https://doi.org/10.1111/ele.12795

    Liu, X., Yin, X. & Ma, J. (2001) Biology of Marine-Fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing, 860 pp., 82 pls.

    MacGillivray, P.H. (1895) A Monograph of the Tertiary Polyzoa of Victoria. Transactions of the Royal Society of Victoria, 4, 1166.

    https://doi.org/10.5962/bhl.title.6076

    Maturo, F.J.S. & Schopf, T.J.M. (1968) Ectoproct and entoproct type material: re-examination of species from Bermuda and New England collected by A.E. Verrill, J.W. Dawson and E. Desor. Postilla, 120, 195.

    Manzoni, A. (1870) Briozoi fossili italiani. Quarta contribuzione. Sitzungsberichte der Akademie der Wissenschaften in Wien, Abteilung 1, 61, 323–349.

    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

    Milne-Edwards, H. (1838) Mémoire sur les Polypes du genre des Tubulipores. Annales des Sciences naturelles, Zoologie & Biologie animale, 8, 321 –338.

    Ngai, N.D., Cu, N.D. & Tuyet, A. (2013) Coral degradation and ability of rehabilitation of coral reefs in Co To Archipelago, Quang Ninh province, Vietnam. Deep-Sea Research II, 96, 50–55.

    https://doi.org/10.1016/j.dsr2.2013.04.017

    Neviani, A. (1895) Briozoi fossili della Farnesina e Monte Mario presso Roma. Palaeontographia Italica, 1, 77–139, pls. 5–6.

    Norman, A.M. (1903a) Notes on the natural history of East Finmark. Polyzoa. Annals and Magazine of Natural History, Series 7, 11, 567–598, pl. 13.

    https://doi.org/10.1080/00222930308678818

    Norman, A.M. (1903b) Notes on the natural history of East Finmark. Polyzoa. Annals and Magazine of Natural History, Series 7, 12, 87–128.

    https://doi.org/10.1080/00222930308678831

    O’Dea, A. & Okamura, B. (1999) Influence of seasonal variation in temperature, salinity and food availability on module size and colony growth of the estuarine bryozoan Conopeum seurati. Marine Biology, 135, 581–588.

    https://doi.org/10.1007/s002270050659

    O’Dea, A. & Okamura, B. (2000) Intracolony variation in zooid size in cheilostome bryozoans as a new technique for investigation palaeoseasonality. Palaeogeography Palaeoclimatology Palaeoecology, 162, 319–332.

    https://doi.org/10.1016/S0031-0182(00)00136-X

    Osburn, R.C. (1952) Bryozoa of the Pacific Coast of North America. Part 2. Cheilostomata-Ascophora. Allan Hancock Pacific Expeditions, 14, 271–611.

    Ostrovsky, A.N. (2013) Evolution of Sexual Reproduction in Marine Invertebrates: Example of Gymnolaemate Bryozoans. Springer, Dordrecht, 356 pp.

    https://doi.org/10.1007/978-94-007-7146-8

    Pallas, P.S. (1766) Elenchus Zoophytorum Sistens Generum Adumbrationes Generaliores et Speciarum Cognitarum Cuccintas Descriptiones cum Selectis Auctorus Synonymis. Petrum van Cleef, Hagae-Comitum, 451 pp.

    https://doi.org/10.5962/bhl.title.6595

    Parker, W.K. & Jones, T.R. (1860) On the nomenclature of the Foraminifera. Annals and Magazine of Natural History, Series 3, 5, 285–297.

    https://doi.org/10.1080/00222936008697218

    Pergens, E. & Meunier, A. (1887) La faune des Bryozoaires garumniens de Faxe. Annales de la Société Royale Malacologique de Belgique, Memoires, 21, 187–242.

    Richardson-White, S. & Walker, S.E. (2011) Diversity, taphonomy and behavior of encrusting foraminifera on experimental shells deployed along a shelf-to-slope bathymetric gradient, Lee Stocking Island, Bahamas. Palaeogeography Palaeoclimatology Palaeoecology, 312, 305–324.

    https://doi.org/10.1016/j.palaeo.2011.02.021

    Ristedt, H. & Hillmer, G. (1985) The cheilostomate bryozoan fauna from shallow waters of the Hilutangan Channel, Cebu, Philippines: part 1. The Philippine Scientist, 22, 133–143.

    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

    Scholz, J. (1991) Die Bryozoanfauna der philippinischen Riffregion Cebu. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg, 71, 253–403.

    Scholz J, & Cusi, M.A.V. (1991) Paleoecologic implications of modern coral and bryozoan communities from southern Leyte, Philippines. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg, 71, 405–431.

    Seo, J.E. (1994) Two species of Celleporaria (Cheilostomata: Bryozoa) from Korea. Korean Journal of Systematic Zoology, 10, 189–197.

    Skinner, L.F. (2014) Foraminifera as predators on encrusting bryozoans. Journal of Foraminiferal Research, 44, 58–61.

    https://doi.org/10.2113/gsjfr.44.1.58

    Smitt, F.A. (1867) Kritisk förteckning öfver Skandinaviens Hafs-Bryozoer II. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 23, 395–534, pls. 3–13.

    Smitt, F.A. (1868) Kritisk förteckning öfver Skandinaviens Hafs-Bryozoer, IV. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 25, 3–320.

    Smitt, F.A. (1873) Floridan Bryozoa, collected by Count L. F. de Pourtales. Part 2. Kongliga Svenska Vetenskaps-Akademiens Handlingar, 11, 1–83, pls. 1–13.

    Soule, D.F. & Soule, J.D. (1970) New species of Thalamoporella (Ectoprocta) from Hawaii, examined by scanning electron microscopy. American Museum Novitates, 2417, 1–18.

    Soule, D.F., Soule, J.D. & Chaney, H.W. (1991a) Some little-known genera of Thalamoporellidae: Thairopora, Diploporella, and new genera Marsupioporella and Thalamotreptos. In: Bigey, F.P. & d’Hondt, J.-L. (Eds.), Bryozoaires Actuels et Fossiles: Bryozoa Living and Fossil. Société des Sciences Naturelles de l’Ouest de la France, Nantes, pp. 447–464.

    Soule, D.F., Soule, J.D. & Chaney, H.W. (1991b) New tropical Pacific and Indian Ocean Cleidochasmatidae (Cheilostomata: Ascophora). In: Bigey, F.P. & d’Hondt, J.-L. (Eds.), Bryozoaires Actuels et Fossiles: Bryozoa Living and Fossil. Société des Sciences Naturelles de l’Ouest de la France, Nantes, pp. 465–486.

    Soule, D.F., Soule, J.D. & Chaney, H.W. (1992) The genus Thalamoporella worldwide (Bryozoa Anasca): morphology, evolution and speciation. Irene McCulloch Foundation Monograph Series, 1, 1–93.

    Soule, D.F., Soule, J.D. & Chaney, H.W. (1995) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and western Santa Barbara channel. Irene McCulloch Foundation Monograph Series, 2, 1–344.

    Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña, Z.A., Finlayson, N., Halpern, B.S., Jorge, M.A., Lombana, A., Lourie, S.A., Martin, K.D., McManus, E., Molnar, J., Recchia, C.A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57, 573–583.

    https://doi.org/10.1641/B570707

    Suwa, T. & Mawatari, S.F. (1998) Revision of seven species of Microporella (Bryozoa, Cheilostomatida) from Hokkaido, Japan, using new taxonomic characters. Journal of Natural History, 32, 895–922.

    https://doi.org/10.1080/00222939800770461

    Taylor, P.D. (2016) Competition between encrusters on marine hard substrates and its fossil record. Paleontology, 59, 481–497.

    https://doi.org/10.1111/pala.12239

    Taylor, P.D. & Tan, S.-H.A. (2015) Cheilostome Bryozoa from Penang and Langkawi, Malaysia. European Journal of Taxonomy, 149, 1–34.

    https://doi.org/10.5852/ejt.2015.149

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

    Tilbrook, K.J. (1998) The species of Antropora Norman, 1903 (Bryozoa: Cheilostomatida), with the description of a new genus in the Calloporoidea. Records of the South Australian Museum, 31, 25–49.

    Tilbrook, K.J. (1999) Description of Hippopodina feegeensis and three other species of Hippopodina Levinsen, 1909 (Bryozoa: Cheilostomatida). Journal of the Zoological Society of London, 247, 449–456.

    https://doi.org/10.1111/j.1469-7998.1999.tb01008.x

    Tilbrook, K.J. (2006) Cheilostomatous Bryozoa of the Solomon Islands. Santa Barbara Museum of Natural History, Santa Barbara, 386 pp.

    Tilbrook, K.J. (2012) Review of the bryozoan genus Bryopesanser Tilbrook, 2006 (Escharinidae: Cheilostomata) with the description of 11 new species. Zootaxa, 3165 (1), 39–63.

    https://doi.org/10.11646/zootaxa.3165.1.3

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

    https://doi.org/10.1111/j.1096-3642.2001.tb01309.x

    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.

    Walker, S.E., Parsons-Hubbard, K., Richardson-White, S., Brett, C. & Powell, E. (2011) Alpha and beta diversity of encrusting foraminifera that recruit to long-term experiments along a carbonate platform-to-slope gradient: paleoecological and paleoenvironmental implications. Palaeogeography Palaeoclimatology Palaeoecology, 312, 325–349.

    https://doi.org/10.1016/j.palaeo.2011.04.028

    Waters, A.W. (1887) On Tertiary cyclostomatous Bryozoa from New Zealand. Quarterly Journal of the Geological Society, London, 43, 337–350, pl. 18.

    https://doi.org/10.1144/GSL.JGS.1887.043.01-04.27

    Winston, J.E. (2005) Re-description and revision of Smitt’s “Floridan Bryozoa” in the collection of the Museum of Comparative Zoology, Harvard University. Virginia Museum of Natural History Memoir, 7, i–ix + 1–147.

    Winston, J.E. (2012) Dispersal in marine organisms without a pelagic larval phase. Integrative and Comparative Biology, 52, 447–457.

    https://doi.org/10.1093/icb/ics040

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

    Winston, J.E. & Woollacott, R.M. (2009) Scientific results of the Hassler expedition. Bryozoa. No. 1. Barbados. Bulletin of the Museum of Comparative Zoology, 159, 239–300.

    https://doi.org/10.3099/0027-4100-159.5.239

    Woodin, S.A. & Jackson, J.B.C. (1979) Interphyletic competition among marine benthos. Integrative and Comparative Biology, 19, 1029–1043.

    https://doi.org/10.1093/icb/19.4.1029

    Yang, J.Y., 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