Abstract
This work describes Bryozoa of the orders Cyclostomata and Ctenostomata found associated with polymetallic nodules collected by box-coring in the eastern part of the Russian exploration area of the Clarion-Clipperton Fracture Zone (CCFZ) under contract to Yuzhmorgeologiya. Scanning electron microscopic study of 358 cyclostome colonies and 14 ctenostome colonies from 4510–5280 m depth has resulted in the recognition of two new species of Ctenostomata, and 14 new species, nine new genera and two new families of Cyclostomata; three additional species of Cyclostomata are left in open nomenclature pending the discovery of missing reproductive characters. The taxonomic novelty is thus notable. One of the new Ctenostomata represents the first living example of the previously monotypic Late Cretaceous genus Pierrella. Twelve of the new cyclostome taxa have well-developed gonozooids, indicating that embryonic cloning (polyembryony) is normal in this deep-sea environment. On the other hand, one indeterminate tubuliporine and two rectangulates have dimorphic peristomes. In the latter two cases, enough mature colonies were found to suggest that this feature is normal, and that the dimorphic zooids are possibly female—in other words, capacious incubation chambers are apparently lacking, and therefore polyembryony would also be lacking or reduced. In one of these species, evidence is presented to suggest that the ancestrular zooid can reproduce precociously. Of the species reported here, only one has previously been found outside the exploration area, highlighting both the limited knowledge we have of bryozoans in the deep Pacific and/or a fauna that is largely endemic to the nodule environment. An additional 31 species of Cheilostomata have also been discovered that will be described in a subsequent publication. Most bryozoans are macrofaunal-sized, so are both inadequately determinable and overlooked in images obtained by remotely operated vehicles; yet, with 50 species, Bryozoa is the most speciose sessile macrofaunal phylum on the nodules. Nodules constitute hard substrata in an area otherwise mostly inhospitable for Bryozoa, hence mining would lead to loss of critical habitat. Further, as suspension-feeders, bryozoans are highly susceptible to smothering by suspended sediment, and non-mined areas closely adjacent to extraction zones would likely also be affected and their associated bryozoan fauna obliterated. More data are required on the distribution of the CCFZ bryozoan species elsewhere in the east Central Pacific to determine if mining would lead to local taxon extirpation or global extinction at both low and high taxonomic levels.
References
Allman, G.J. (1856) A Monograph of the Freshwater Polyzoa, including all the known species, both British and foreign. The Ray Society, London. viii + 119 pp., 11 pls.
Amon, D.J. (2016) Megafauna and minerals on the Pacific abyss. Deep Sea News, 18 January 2016. Available from: http://www.deepseanews.com/2016/01/megafauna-and-minerals-on-the-pacific-abyss/ (accessed 3 August 2018)
Amon, D.J., Ziegler, A.F., Drazen, J.C., Grischenko, A.V., Leitner, A.B., Lindsay, D.J., Voight, J.R., Wicksten, M.K., Young, C.M. & Smith, C.R. (2017) Megafauna of the UKSRL exploration contract area and eastern Clarion-Clipperton Zone in the Pacific Ocean: Annelida, Arthropoda, Bryozoa, Chordata, Ctenophora, Mollusca. Biodiversity Data Journal, 5, e14598.
https://doi.org/10.3897/BDJ.5.e14598Batson, P.B., Taylor, P.D. & Smith, A.M. (2018) Early astogeny in Hornera (Bryozoa; Cyclostomata; Cancellata). Memoirs of the Association of Australasian Palaeontologists. [in press]
Boardman, R.S., McKinney, F.K. & Taylor, P.D. (1992) Morphology, anatomy, and systematics of the Cinctiporidae, new family (Bryozoa: Stenolaemata). Smithsonian Contributions to Paleobiology, 70, i–iv + 1–81.
https://doi.org/10.5479/si.00810266.70.1Bock, P.E. & Gordon, D.P. (2013) Phylum Bryozoa Ehrenberg, 1831. In: Zhang, Z.-Q. (Ed.), Animal Biodiversity: An outline of higher-level classification and survey of taxonomic richness (Addenda 2013). Zootaxa, 3703, 67–74.
Boetius, A. & Haeckel, M. (2018) Mind the seafloor. Science, 359, 34–36.
https://doi.org/10.1126/science.aap7301Borg, F. (1926) Studies on Recent cyclostomatous Bryozoa. Zoologiska Bidrag från Uppsala, 10, 181–507, 14 pls.
Borg, F. (1944) The stenolaematous Bryozoa. Further Zoological Results of the Swedish Antarctic Expedition 1901–1903, 3 (5), 1–76, 16 pls.
Bronn, H.G. (1825) System der urweltlichen Pflanzenthiere, durch Diagnose, Analyse und Abbildung der Geschlechter erlautert. J.C.B. Mohr, Heidelberg, iv + 47 pp., 7 pls.
Brown, J.H., Gillooly, J.F., Allen, A.P., Savage, V.M. & West, G.B. (2004) Toward a metabolic theory of ecology. Ecology, 8, 1771–1789.
https://doi.org/10.1890/03-9000Busk, 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. W. Boone, London, pp. 343–402, pl. 1.
Busk, G. (1884) Report on the Polyzoa collected by H.M.S. Challenger during the years 1873–76. Part I.—The Cheilostomata. Report on the Scientific Results of the Voyage of H.M.S. Challenger, Zoology, 10 (30), i–xxiv + 1–216, 36 pls.
Busk, G. (1886) Report on the Polyzoa collected by H.M.S. Challenger during the years 1873–76. Part II.—The Cyclostomata, Ctenostomata, and Pedicellinea. Report on the Scientific Results of the Voyage of H.M.S. Challenger, Zoology, 17 (50), i–viii + 1–47, 10 pls.
Canu, F. (1916) [1918] Les ovicelles des Bryozoaires cyclostomes. Bulletin de la Société Géologique de France, Series 4, 16, 324–335.
Costello, M.J. & Chaudhary, C. (2017) Marine biodiversity, biogeography, deep-sea gradients, and conservation. Current Biology, 27, 511–527. [R511–R527]
https://doi.org/10.1016/j.cub.2017.04.060Craw, A. (2013) Deep Seabed Mining: An urgent wake-up call to protect our oceans. Greenpeace, Amsterdam, 19 pp.
Danovaro, R., Snelgrove, P.V.R. & Tyler, P. (2014) Challenging the paradigms of deep-sea ecology. Trends in Ecology and Evolution, 29, 465–475.
David, L. & Pouyet, S. (1986) Bryozoaires abyssaux des campagnes Safari (océan Indien). Annals de l’Institut Océanographique, New Series, 62, 141–191.
Defrance, J.L.M. (1823) Lichenopore. Dictionnaire des Sciences Naturelles, 26, 256–257.
De Smet, B., Pape, E., Riehl, T., Bonifácio, P., Colson, L. & Vanreusel, A. (2017) The community structure of deep-sea macrofauna associated with polymetallic nodules in the eastern part of the Clarion-Clipperton Fracture Zone. Frontiers in Marine Science, 4 (103), 1–14.
https://doi.org/10.3389/fmars.2017.00103Edwards, H.M. (1838) Mémoire sur les Polypes du genre des Tubulipores. Annales des Sciences Naturelles, Series 2, Zoologie 8, 321–338.
Eilertsen, M.H. & Malaquias, M.A.E. (2015) Speciation in the dark: diversification and biogeography of the deep-sea gastropod genus Scaphander in the Atlantic Ocean. Journal of Biogeography, 42, 843–855.
https://doi.org/10.1111/jbi.12471Gordon, D.P. (1987) The deep-sea Bryozoa of the New Zealand region. In: Ross, J.R.P. (Ed.), Bryozoa: Present and Past. Western Washington University, Bellingham, pp. 97–104.
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.6.243Gordon, D.P. & Taylor, P.D. (2010) New seamount- and ridge-associated cyclostome Bryozoa from New Zealand. Zootaxa, 2533, 43–68.
Gray, J.E. (1848) List of the Specimens of British Animals in the Collection of the British Museum. Part 1. Centroniae or radiated animals. Trustees of the British Museum (natural History), London, xiii + 173 pp.
Gregory, J.W. (1896) Catalogue of the Fossil Bryozoa in the Department of Geology, British Museum (Natural History). The Jurassic Bryozoa. Trustees of the British Museum (Natural History), London, 239 pp., 11 pls.
Gregory, J.W. (1899) Catalogue of the Fossil Bryozoa in the Department of Geology, British Museum (Natural History). The Cretaceous Bryozoa. Vol. I. Trustees of the British Museum (Natural History), London, xiv + 457 pp., 17 pls.
Grischenko, A.V. (2013) First record of a bathyal bryozoan fauna from the Sea of Japan. Deep-Sea Research II, 86–87, 172–180.
https://doi.org/10.1016/j.dsr2.2012.08.003Grischenko, A.V. & Chernyshev, A.V. (2015) Triticella minini – a new ctenostome bryozoan from the abyssal plain adjacent to the Kuril-Kamchatka Trench. Deep-Sea Research II, 111, 343–350.
https://doi.org/10.1016/j.dsr2.2014.08.004Grischenko, A.V. & Chernyshev, A.V. (2017) Deep-water Bryozoa from the Kuril Basin, Sea of Okhotsk. Deep-Sea Research II, 113.
https://doi.org/10.1016/j.dsr2.2017.09.011Harmelin, J.-G. & Hondt, J.-L. (1982) Bryozoaires cyclostomes bathyaux des campagnes océanographiques de l’“Atlantis II”, du “Chain” et du “Knorr” (1967–1972). Bulletin du Muséum National d’Histoire Naturelle, 4A, 4, 3–23.
Hayward, P.J. (1978) Two new species of Ctenostomata (Bryozoa) from the Norwegian Sea. Sarsia, 63, 159–162.
https://doi.org/10.1080/00364827.1978.10411334Hayward, P.J. (1981) The Cheilostomata (Bryozoa) of the deep sea. Galathea Report, 15, 21–68, pl. 5.
Hayward, P.J. & Ryland, J.S. (1985) Cyclostome bryozoans. Synopses of the British Fauna, New Series, 34, 1–147.
Hincks, T. (1880) A History of the British Marine Polyzoa. Van Voorst, London, cxli + 601 pp., 83 pls.
Hondt, J.-L. (1982) Les Bryozoaires eurystomes abyssaux. Comptes Rendu des Séances de la Société de Biogéographie, 58, 30–48.
Hondt, J.-L. (1983) Tabular keys for identification of the Recent ctenostomatous Bryozoa. Mémoires de l’Institut Océanographique, Monaco, 14, 1–134.
Hondt, J.-L. & Gordon, D.P. (1996) Bryozoa: Cténostomes et Cheilostomes (Cellularines, Scrupariines et Malacostèges) des Campagnes MUSORSTOM autour de la Nouvelle-Calédonie. In: Crosnier, A. (Ed.), Résultats des Campagnes MUSORSTOM. Vol. 15. Mémoires du Muséum National d'Histoire Naturelle, 168, pp. 55–123.
Hondt, J.-L. & Mawatari, S.F. (1987) Description de deux nouvelles espèces d’Arachnoidea (Bryozoa, Ctenostomida) provenant de la campagne océanographique KAIKO I (Japon). Cahiers de Biologie Marine, 28, 373–380.
Hondt, J.-L. & Schopf, T.J.M. (1984) [1985] Bryozoaires des grandes profondeurs receuillis lors des campagnes océanographiques de la Woods Hole Oceanographic Institution de 1961 à 1968. Bulletin du Muséum National d’Histoire Naturelle, Series 4, 4, 6A, 907–973.
Hughes, D.J. (1987) Gametogenesis and embryonic brooding in the cheilostome bryozoan Celleporella hyalina. Journal of Zoology, 212, 691–711.
https://doi.org/10.1111/j.1469-7998.1987.tb05965.xJablonski, D., Sepkoski, J.J., Bottjer, D.J. & Sheehan, P.M. (1983) Onshore-offshore patterns in the evolution of Phanerozoic shelf communities. Science, 222, 1123–1125.
https://doi.org/10.1126/science.222.4628.1123Jebram, D. (1986) The ontogenetical and supposed phylogenetical fate of the parietal muscles in the Ctenostomata (Bryozoa). Zeitschrift für Zoologische und Systematische Evolutionsforschung, 24, 58–82.
https://doi.org/10.1111/j.1439-0469.1986.tb00616.xJenkins, H.L. & Taylor, P.D. (2014) New or poorly known skeletal features in the early astogeny of some cyclostome bryozoans. Studi Trentini di Scienze Naturali, 94, 125–130.
Jenkins, H.L., Bishop, J.D.D. & Hughes, R.N. (2015) Prudent female allocation by modular hermaphrodites: female investment is promoted by the opportunity to outcross in cyclostome bryozoans. Biological Journal of the Linnean Society, 116, 593–602.
https://doi.org/10.1111/bij.12631Jenkins, H.L., Waeschenbach, A., Okamura, B., Hughes, R.N. & Bishop, J.D.D. (2017) Phylogenetically widespread polyembryony in cyclostome bryozoans and the protracted asynchronous release of clonal brood mates. PLoS ONE, 12 (1), e0170010.
https://doi.org/10.1371/journalpone.0170010Jones, D.O., Kaiser, S., Sweetman, A.K., Smith, C.R., Menot, L., Vink, A., Trueblood, D., Greinert, J., Billett, D.S.M., Arbizu, P.M., Radziejewska, T., Singh, R., Ingole, B., Stratmann, T., Simon-Lledo, E., Durden, J.M. & Clark, M.R. (2017) Biological responses to disturbance from simulated deep-sea polymetallic nodule mining. PLoS ONE, 12 (2), e0171750.
https://doi.org/10.1371/journal.pone.0171750Jullien, J. (1882) Dragages du “Travailleur”, Bryozoaires. Espèces draguées dans l’Océan Atlantique en 1881. Espèces nouvelles ou incomplètement décrites. Extrait du Bulletin de la Société Zoologique de France, 7, 1–33.
Jullien, J. (1885) Monographie des Bryozoaires d'eau douce. Bulletin de la Société Zoologique de France, 10, 91–207.
Kaiser, S., Smith, C.R. & Arbizu, P.M. (2017) Editorial: Biodiversity of the Clarion Clipperton Fracture Zone. Marine Biodiversity, 47, 259–264.
https://doi.org/10.1007/s12526-017-0733-0Lamouroux, 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 premiers appartenant à l'Histoire naturelle des Zoophytes d'Ellis et Solander. V. Agasse, Paris, 115 pp., 84 pls.
Lim, S.-C., Wiklund, H., Glover, A.G., Dahlgren, T.G. & Tan, K.-S. (2017) A new genus and species of abyssal sponge commonly encrusting polymetallic nodules in the Clarion-Clipperton Zone, East Pacific Ocean. Systematics and Biodiversity, 15 (6), 507–519.
https://doi.org/10.1080/14772000.2017.1358218Lindner, A., Cairns, S.D. & Cunningham, C.W. (2008) From offshore to onshore: Multiple origins of shallow-water corals from deep-sea ancestors. PLoS ONE, 3 (6), 32429.
https://doi.org/10.1371/journal.pone.0002429McKinney, M.J. & Dewel, R.A. (2002) The ctenostome collar—an enigmatic structure. In: Wyse Jackson, P.N., Buttler, C.J. & Spencer Jones, M.E. (Eds.), Bryozoan Studies 2001. Swets & Zeitlinger, Lisse, pp. 191–197.
Miljutin, D.M., Miljutina, M.A., Arbizu, P.M. & Galéron, J. (2011) Deep-sea nematode assemblage has not recovered 26 years after experimental mining of polymetallic nodules (Clarion-Clipperton Fracture Zone, tropical eastern Pacific). Deep-Sea Research Part I, 58, 885–897.
https://doi.org/10.1016/j.dsr.2011.06.003Miller, K.A., Thompson, K.F., Johnston, P. & Santillo, D. (2018) An overview of seabed mining including the current state of development, environmental impacts, and knowledge gaps. Frontiers in Marine Science, 4 (418), 1–24.
https://doi.org/10.3389/fmars.2017.00418Moyano, G.H.I. (1991) Bryozoa from deep-sea waters in Chile: Cyclostomata. In: Bigey, F.P. (Ed.), Bryozoaires actuels et fossiles: Bryozoa Living and Fossil. Société des Sciences Naturelles de l’Ouest de la France, Mémoire Hors-Série, 1, 281–290.
Orbigny, A.D. (1851–54) Bryozoaires. Paléontologie française. Description des Mollusques et Rayonnées fossiles. Terrains Crétacés, 5, 1–188 (1851), 185 bis–472 (1852), 473–984 (1853), 985–1192 (1854), pls. 600–800 [fide Sherborn (1899)].
Ostrovsky, A.N. (1998a) The genus Anguisia as a model of a possible origin of erect growth in some Cyclostomatida (Bryozoa). Zoological Journal of the Linnean Society, 124, 355–367.
https://doi.org/10.1111/j.1096-3642.1998.tb00581.xOstrovsky, A.N. (1998b) Comparative studies of ovicell anatomy and reproductive patterns in Cribrilina annulata and Celleporella hyalina (Bryozoa: Cheilostomatida). Acta Zoologica, 79, 287–318.
https://doi.org/10.1111/j.1463-6395.1998.tb01280.xOstrovsky, A.N. (2013) Evolution of Sexual Reproduction in Marine Invertebrates: Example of gymnolaemate Bryozoa. Springer, Dordrecht, 356 pp.
https://doi.org/10.1007/978-94-007-7146-8Ostrovsky, A.N. & Taylor, P.D. (1996) Systematics of some Antarctic Idmidronea and Exidmonea (Bryozoa: Cyclostomata). Journal of Natural History, 30, 1549–1575.
https://doi.org/10.1080/00222939600770881Pergens, E. & Meunier, A. (1886)[1887] La faune des Bryozoaires garumniens de Faxe. Annales de la Société Royale Malacologique de Belgique, 21, 187–242, pls. 9–13.
Radziejewska, T. (2014) Characteristics of the sub-equatorial north-eastern Pacific's ocean abyss, with a particular reference to the Clarion-Clipperton Fracture Zone. In: Radziejewska, T. (Ed.), Meiobenthos in the Sub-equatorial Pacific Abyss: A proxy in anthropogenic impact evaluation. Springer, Dordrecht, pp. 13–28.
Reverter-Gil, Ó., Souto, J. & Fernández-Pulpeiro, E. (2016) Bryozoa I. Ctenostomata. Fauna Iberica, 43, 1–305.
Rex, M.A., McClain, C.R., Johnson, N.A., Etter, R.J., Allen, J.A., Bouchet, P. & Warén, A. (2005) A source-sink hypothesis for abyssal diversity. The American Naturalist, 165, 163–178.
Ross, J.R.P. & Ross, C.A. (1996) Bryozoan evolution and dispersal and Paleozoic sea-level fluctuations. In: Gordon, D.P., Smith, A.M. & Grant-Mackie, J.A. (Eds.), Bryozoans in Space and Time. NIWA, Wellington, pp. 243–258.
Ryland, J. (1996) Polyembryony 'paradox': the case of cyclostomate Bryozoa’. Trends in Ecology and Evolution, 11, 26.
https://doi.org/10.1016/0169-5347(96)80240-0Schäfer, P. (1991) Brutkammern der Stenolaemata (Bryozoa): Konstruktionsmorphologie und phylogenetische Bedeutung. Courier Forschungsinstitut Senckenberg, 136, 1–263.
Sherborn, C.D. (1899) On the dates of the "Paléontologie Française" of d'Orbigny. Geological Magazine, Series 4, 6, 223–225.
Schopf, T.J.M. (1969) Geographic and depth distribution of the phylum Ectoprocta from 200 to 6,000 meters. Proceedings of the American Philosophical Society, 113, 464–474.
Schwaha, T., Handschuh, S., Ostrovsky, A.N. & Wanninger, A. (2018) Morphology of the bryozoan Cinctipora elegans (Cyclostomata, Cinctiporidae) with the first data on its sexual reproduction and the cyclostome neuro-muscular system. BMC Evolutionary Biology, 18, 92.
https://doi.org/10.1186/s12862-018-1206-1Silén, L. & Harmelin, J.-G. (1974) Observations on living Diastoporidae (Bryozoa Cyclostomata), with special regard to polymorphism. Acta Zoologica, 55, 81–96.
https://doi.org/10.1111/j.1463-6395.1974.tb00182.xSmitt, F.A. (1867) Kritisk förteckning öfver Skandinaviens Hafs-Bryozoer. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 23, 395–533, pls. 3–13.
Taylor, P.D. (1993) Bryozoa. In: Benton, M.J. (Ed.), The Fossil Record. Chapman & Hall, London, pp. 465–489.
Taylor, P.D. & Grischenko, A.V. (1999) Rodinopora gen. nov. and the taxonomy of fungiform cyclostome bryozoans. Species Diversity, 4, 9–33.
https://doi.org/10.12782/specdiv.4.9Taylor, P.D. & Sequeiros, L. (1982) Toarcian bryozoans from Belchite in north-east Spain. Bulletin of the British Museum (Natural History), Geology, 37, 117–129.
Taylor, P.D. & Weedon, M. (2000) Skeletal ultrastructure and phylogeny of cyclostome bryozoans. Zoological Journal of the Linnean Society, 128, 337–399.
https://doi.org/10.1111/j.1096-3642.2000.tb01521.xTaylor, P.D. & Wilson, M.A. (1999) Middle Jurassic bryozoans from the Carmel Formation of southwestern Utah. Journal of Paleontology, 73, 816–830.
https://doi.org/10.1017/S0022336000040671Taylor, P.D. & Zatón, M. (2008) Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos (Cyclostomata: Oncousoeciidae). Journal of Natural History, 42, 2557–2574.
https://doi.org/10.1080/00222930802277640Thuy, B., Gale, A.S., Kroh, A., Kucera, M., Numberger-Thuy, Reich, M. & Stöhr, S. (2012) Ancient origin of the modern deep-sea fauna. PLoS ONE, 7 (10), e46913.
https://doi.org/10.1371/journal.pone.0046913Van Dover, C.L., Ardron, J.A., Escobar, E., Gianni, M., Gjerde, K.M., Jaeckel, A., Jones, D.O.B., Levin, L.A., Niner, H.J., Pendleton, L., Smith, C.R., Thiele, T., Turner, P.J., Watling, L. & Weaver, P.P.E. (2017) Biodiversity loss from deep-sea mining. Nature Geoscience, 10, 464–465.
https://doi.org/10.1038/ngeo2983Vanreusel, A., Hilario, A., Ribeiro, P.A., Menot, L. & Arbizu, P.M. (2016) Threatened by mining, polymetallic nodules are required to preserve abyssal fauna. Scientific Reports, 6 (26808), 1–6.
Voigt, E. (1985) The Bryozoa of the Cretaceous - Tertiary boundary. In: Nielsen, C. & Larwood, G.P. (Eds.), Bryozoa: Ordovician to Recent. Olsen & Olsen, Fredensborg, pp. 329–342.
Waters, A.W. (1887) On Tertiary cyclostomatous Bryozoa from New Zealand. Quarterly Journal of the Geological Society of London, 43, 337–350, pl. 18.
https://doi.org/10.1144/GSL.JGS.1887.043.01-04.27Waters, A.W. (1889) 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 H.M.S. Challenger, Zoology, 31 (79), 1–41, 3 pls.
Wiklund, H., Taylor, J.D., Dahlgren, T.G., Todt, C., Ikebe, C., Rabone, M. & Glover, A.G. (2017) Abyssal fauna of the UK-1 polymetallic nodule exploration area, Clarion-Clipperton Zone, central Pacific Ocean: Mollusca. Zookeys, 707, 1–46.
https://doi.org/10.3897/zookeys.707.13042Wilson, M.A. & Taylor, P.D. (2013) Palaeoecology, preservation and taxonomy of encrusting ctenostome bryozoans inhabiting ammonite body chambers in the Late Cretaceous Pierre Shale of Wyoming and South Dakota. In: Ernst, A., Schäfer, P. & Scholz, J. (Eds.), Bryozoan Studies 2010. Lecture Notes in Earth System Sciences, 143, pp. 419–433.
https://doi.org/10.1007/978-3-642-16411-8_28