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The zoological dry collection of the Swedish Museum of Natural History in Stockholm includes an important, historical bryozoan section that is rich in species and specimens and also diverse from a geographical point of view. This collection also contains the type specimens of the type species of some cheilostome bryozoan genera introduced by several naturalists and bryozoologists between the mid-1800s and 1900s. With a few exceptions, these have not been revised since the advent of scanning electron microscopy as a standard tool for bryozoan taxonomy. Here, the type specimen(s) of the type species of the following six cheilostome genera are described and illustrated using SEM micrographs for the first time: Cheilopora Levinsen, 1909; Fedorella Silén, 1947; Floridina Jullien, 1882; Lepraliella Levinsen, 1917; Smittipora Jullien, 1882; and Stenopsella Bassler, 1952. The type specimen(s) of the type species of the recently introduced Terwasipora Reverter-Gil & Souto, 2019 and the relatively recently revised Doryporella Norman, 1903 are also illustrated for the first time. This revision has identified some erroneous geographical records for some of the species/genera examined, and has led to the proposed synonymy of Stenopsella with Gigantopora Ridley, 1881. Lectotypes have also been selected. All of the images produced will also be publicly available through the SMNH online catalogue. The digitisation of natural history museum collections, with prioritisation of historical type specimens, is of paramount importance to facilitate access to the fundamental taxonomic units for scientists worldwide.

Genus Terwasipora Reverter-Gil & Souto, 2019 Type species Lepralia complanata Norman, 1864, Published as part of Martino, Emanuela Di, 2022, Revision of the type species of some cheilostome bryozoan genera in the collection of the Swedish Museum of Natural History, pp. 157-181 in Zootaxa 5125 (2) on page 175, DOI: 10.11646/zootaxa.5125.2.4, http://zenodo.org/record/6424502, {"references":["Reverter-Gil, O. & Souto, J. (2019) Watersiporidae (Bryozoa) in Iberian waters: an update on alien and native species. Marine Biodiversity, 49, 2735 - 2752. https: // doi. org / 10.1007 / s 12526 - 019 - 01003 - 4","Norman, A. M. (1864) On undescribed British Hydrozoa, Actinozoa and Polyzoa. Annals and Magazine of Natural History, 13, 82 - 90. https: // doi. org / 10.1080 / 00222936408681578"]}

Terwasipora complanata (Norman, 1864) (Fig. 10) Lepralia complanata Norman, 1864: 85, pl. 10, fig. 4. Terwasipora complanata: Reverter-Gil & Souto 2019: 2743, figs 5, 6. Material examined. Holotype by monotypy SMNH-Type-1772, North Atlantic Ocean, Ireland, encrusting a rock. Leg. unknown. Remarks. The description of this species is omitted because accurate descriptions are available in several papers, some very recent (e.g. Reverter-Gil & Souto 2019, Ramalho et al. 2022). However, the original material described by Norman (1864) has never been described and/or illustrated. The syntype SMNH-Type-1772 (Fig. 10A) consists of a fan-shaped colony, about 5.8 x 8.6 mm, encrusting a granitic rock, about 9 x 11.5 cm, sharing the substrate with spirorbid polychaetes. Norman (1864) acknowledged having found the specimen in the collection of British Zoophytes of his late friend Mr. Barlee that he inherited. This information agrees with the label (Fig. 10B), as does the description of the substrate provided by Norman (1864) as coarse-grained granite. While Norman (1864) claimed not to have detailed information about the collecting locality, apart from being British, the label reports Ireland as the probable origin, which seems plausible given that the geology of the northern coast of Ireland is characterized by igneous rocks and granitic intrusions. This species has a wide Atlanto-Mediterranean distribution. Confirmed records are from the Isles of Scilly (off Cornwall, England) encrusting shells and rocks (Hayward & Ryland 1999), several localities along the Atlantic Iberian coast (Reverter-Gil & Souto 2019), the Strait of Gibraltar (e.g. López de la Cuadra 1991), the Alboran Sea (Ramalho et al. 2022), and the western Mediterranean (Gautier 1962, Novosel & Pozar-Domac, 2001, Rosso & Di Martino 2016)., Published as part of Martino, Emanuela Di, 2022, Revision of the type species of some cheilostome bryozoan genera in the collection of the Swedish Museum of Natural History, pp. 157-181 in Zootaxa 5125 (2) on pages 175-176, DOI: 10.11646/zootaxa.5125.2.4, http://zenodo.org/record/6424502, {"references":["Norman, A. M. (1864) On undescribed British Hydrozoa, Actinozoa and Polyzoa. Annals and Magazine of Natural History, 13, 82 - 90. https: // doi. org / 10.1080 / 00222936408681578","Reverter-Gil, O. & Souto, J. (2019) Watersiporidae (Bryozoa) in Iberian waters: an update on alien and native species. Marine Biodiversity, 49, 2735 - 2752. https: // doi. org / 10.1007 / s 12526 - 019 - 01003 - 4","Ramalho, L. V., Rodrigues-Aporta, R. & Gofas, S. (2022) Preliminary account on the bryozoans of the Alboran platform (Western Mediterranean), with description of two new species. Zootaxa, 5094 (1), 53 - 91. https: // doi. org / 10.11646 / zootaxa. 5094.1.2","Hayward, P. J. & Ryland, J. S. (1999) Cheilostomatous Bryozoa. Part 2. Hippothoidea - Celleporoidea. Synopses of the British Fauna. New Series. Vol. 14. Field Studies Council, Shrewsbury, 416 pp.","Lopez de la Cuadra, C. M. (1991) Estudio sistematico de los Briozoos Queilostomados (Bryozoa: Cheilostomida) del Estrecho de Gibraltar y areas proximas. PhD Thesis. Universidad de Sevilla, Sevilla, 239 pp.","Gautier, Y. V. (1962) Recherches ecologiques sur les Bryozoaires chilostomes en Mediterranee occidentale. Recueil des travaux de la Station marine d'Endoume, 38, 1 - 434.","Novosel, M. & Pozar-Domac, A. (2001) Checklist of Bryozoa of the Eastern Adriatic Sea. Natura Croatica, 10, 367 - 421.","Rosso, A. & Di Martino, E. (2016) Bryozoan diversity in the Mediterranean Sea: an update. Mediterranean Marine Science, 17, 567 - 607. https: // doi. org / 10.12681 / mms. 1706"]}

Genus Terwasipora Reverter-Gil & Souto, 2019 Type species Lepralia complanata Norman, 1864

Terwasipora complanata (Norman, 1864) (Fig. 12) Lepralia complanata Norman, 1864: 85, pl. 10, fig. 4. Watersipora complanata: Zabala 1986: 395, fig. 128; Hayward & Ryland 1999: 192, figs 74b, 75; Reverter-Gil & Fern��ndez- Pulpeiro 2001: 122; De Blauwe 2009: 372, figs 397, 398; Reverter-Gil et al. 2014: 26. Terwasipora complanata: Reverter-Gil & Souto 2019: 2743, figs 5, 6, 8. Material examined. MNCN 25.03/4282: BV12, 112��� 120 m; UMA coll.; one dead fragment on shell; MNCN 25.03/4283: BV12, 112��� 120 m; UMA coll.; four living fragments on shell and three loose dead fragments. Remarks. Several specimens from the Mediterranean Sea assigned to T. complanata were reviewed by Reverter- Gil & Souto (2019), who maintained as valid the records of this species from Catalonia and the Strait of Gibraltar (Zabala 1986; L��pez de la Cuadra & Garc��a-G��mez 1988; L��pez de la Cuadra 1991; see Reverter-Gil & Souto 2019 for comments about its geographic distribution). The Alboran colonies have slightly smaller zooids (L 600���800 ��m x W 350���600 ��m) but within the range reported by Reverter-Gil & Souto (2019)., Published as part of Ramalho, La��s V., Rodr��guez-Aporta, Raquel & Gofas, Serge, 2022, Preliminary account on the bryozoans of the Alboran platform (Western Mediterranean), with description of two new species, pp. 53-91 in Zootaxa 5094 (1) on pages 76-77, DOI: 10.11646/zootaxa.5094.1.2, http://zenodo.org/record/5964865, {"references":["Norman, A. M. (1864) On undescribed British Hydrozoa, Actinozoa and Polyzoa. Annals and Magazine of Natural History, Series 3, 13 (73), 82 - 90. https: // doi. org / 10.1080 / 00222936408681578","Zabala, M. (1986) Fauna dels Briozous dels Paisos Catalans. Institut d'Estudis Catalans, Arxius de la Seccio de Ciencies, Barcelona, 836 pp.","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. The Linnean Society of London, London, pp. 1 - 416.","Reverter-Gil, O., Souto, J. & Fernandez-Pulpeiro, E. (2014) Annotated checklist of Recent marine Bryozoa from continental Portugal. Nova Acta Cientifica Compostelana, Bioloxia, 21, 1 - 55.","Reverter-Gil, O. & Souto, J. (2019) Watersiporidae (Bryozoa) in Iberian waters: an update on alien and native species. Marine Biodiversity Records, 49, 2735 - 2752. https: // doi. org / 10.1007 / s 12526 - 019 - 01003 - 4","Hassall, A. H. (1841) Supplement to a catalogue of Irish Zoophytes. Annals and Magazine of Natural History, Series 1, 7 (45), 363 - 373. https: // doi. org / 10.1080 / 03745484109442710","Lopez de la Cuadra, C. M. & Garcia-Gomez, J. C. (1988) Briozoos queilostomados del Estrecho de Gibraltar y areas proximas. Cahiers de Biologie Marine, 29, 21 - 36."]}

The Alboran platform, located in the Alboran Sea, is an area of high productivity due to the mixture of the Atlantic and Mediterranean waters, which produce an almost permanent upwelling in the region. It has been considered one of the main biodiversity hotspots in the Mediterranean Sea. Previous taxonomic studies on bryozoans carried out around the Alboran platform mentioned a total of 127 species collected at depths between 28 and 480 m. Here we report on four Indemares (Inventario y designación de la Red Natura 2000 en áreas marinas del Estado español) samples collected in 2011 and 2012 at depths between 95 and 120 m from a biogenic gravel bottom (i.e. small rocks and shells). Sixty-eight taxa belonging to 52 genera and 34 families were identified. Two species are proposed as new (Escharella similis n. sp. and Hemicyclopora admirabilis n. sp.), eight taxa were identified only to genus level although two of them (Chorizopora sp., Hippothoa sp.) are likely to be undescribed species, and one taxon only at family level. Among the species identified, 30 are new records for the Alboran platform, 13 are new records for the Alboran Sea, and Microporella funbio is recorded for the first time in the Mediterranean Sea. These results covered only a small part of the material sampled (4 out of 44 samples), which suggests that the number of bryozoan species in the Alboran platform is even higher, and also shows the importance of this area for “bryodiversity”.

We report the introduction of the encrusting bryozoan Watersipora subtorquata to Atlantic coasts of Europe. This species is highly invasive, having become common on coastlines throughout cool-temperate areas of the world since the 1980s. Confusion exists over the identity of this and other Watersipora species, which lack characters that are conventionally used in bryozoan systematics. W. subtorquata has not been well distinguished from W. cucullata which, reports dating back to the mid 1800s suggest, is native to the Mediterranean Basin or represents an early shipping introduction. W. cucullata has been placed in synonymy with W. subovoidea, a taxon lacking a holotype. We designate a neotype for W. subovoidea, recognizing its conspecificity with W. cucullata, and demonstrate a simple morphometric means of separating this species from W. subtorquata using zooid feature ratios (operculum area versus total frontal shield). An orange watersiporid population that was first recognized in Guernsey, in the European-Atlantic, in 2007, is shown by morphometric and mitochondrial genetic analysis to match W. subtorquata. It contains the commonest, widely introduced COI haplotype that, along with other evidence, suggests recent transfer via shipping traffic to Europe. A second population (previously referred alternatively to W. aterrima or W. subovoidea) has been reported from Brittany and Bordeaux (Atlantic coastline, France). This population is also aligned with W. subtorquata based on morphometrics and COI haplotype. In contrast to the Guernsey introduction, the earlier French-Atlantic introduction appears related to oyster imports from Japan.

Watersipora subtorquata (d'Orbigny, 1852) (Fig. 5 A–C; Table 6) Cellepora subtorquata d’Orbigny, 1852: 399. Watersipora subtorquata: d’Hondt 1988: 199, figs 6.1– 2; Taylor & Gordon 2002: 536, fig. 1; Chimenz Gusso et al. 2004: 101, figs 9–13; Abdel Salam & Ramadan 2008: 9, fig. 3; Vieira et al. 2014: 155, figs 1–5, 12 – 16, 18–24, 67, 70. Non Watersipora subtorquata: Gordon 1989: 40, pl. 20 B–H; Gordon & Mawatari 1992: 31, fig. 8 E; Ryland et al. 2009: 55, figs 1, 3, 4 A, B, E, F, 5. (More complete synonymy in Vieira et al. 2014). Watersipora subovoidea: Ryland et al. 2009: 54, figs 4 C, D, G, H. Non Cellepora subovoidea d’Orbigny, 1852: 402 (nomen dubium). Material examined. Specimens from Lebanon: 1) Stn 2 A, 3 colonies on rock fragment and pottery; 2) Stn 5 A, small colonies on concretions from cavities; 3) Stn 6 A, 1 colony on Dictyopteris; 4) Stn 6 C, 2 colonies on coralline alga; 5) Stn 7 A, 5 colonies on Pinctada shell; 6) Stn 8 A, 2 colonies; 7) Stn 11 A, 1 small colony on rocky fragment; 8) Stn 12 C, several colonies on rocks; 9) Stn 13 B, several colonies on shells, concretions and pebbles; 10) Stn 14 A, 2 colonies on Spondylus shell; 11) Stn 15 A, several colonies on various substrata. Other material examined: 1) France, Marseille, commercial harbour, Pinede, 3 February 1971, on mussels; same station, 11 December 1982, 6 m, on barnacles; 2) Spain, Tarifa, port, 3 July 1982, 5 m, on Cladocora; 3) Italy, Savona, port, October 1972, 5 m, on barnacles. Description. Colonies small in the collected material, encrusting, unilamellar, moderately pigmented (not observed alive), except for opercula (see below). Zooids subrectangular to hexagonal, relatively short (average L/ W ratio = 1.6) but ranging widely in size (coefficient of variation SD* 100 /X: 14.1% for Az L, 16.1% for Az W). Frontal shield perforated by numerous, evenly distributed, large pseudopores. Orifice distal, large, wider than long (Table 6), with rounded anter developed on about three-quarters of a circle, poster indented by a broad sinus, rounded or sometimes slightly ogival; condyles narrow in frontal view, corresponding to slight bulging of internal arch of anter, superposed to upper side of ‘shoulders’ of sinus, with tip angular or more rounded. Operculum with dark-brown to black thickening in shape of a broad column with slightly arched sides defining a pair of round to oval fenestrae. Tentacles not observed. Remarks. According to the recent revision by Vieira et al. (2014), at least four Watersipora species occur in the Mediterranean Sea. Three of them are relatively common: Watersipora complanata (Norman, 1864), an Atlanto-Mediterranean species lacking a sinus the generic status of which is open to question (Ryland et al. 2009), W. cucullata (Busk, 1854), apparently endemic to the Mediterranean, and W. subtorquata, a non-indigenous species originally described from Brazil but now spread worldwide in tropical and warm-temperate areas (Taylor & Gordon 2002; Vieira et al. 2014). The occurrence in the Mediterranean of the fourth species, W. souleorum Vieira, Spencer Jones & Taylor, 2014, based on type material from the Cape Verde Islands, is attested by specimens collected in the early 20 th century at Naples (Vieira et al. 2014) and recorded by Waters (1909) as Lepralia? cucullata. Therefore, the current presence of this species in the Mediterranean needs to be confirmed. Identification at species-level of Watersipora colonies having a U-shaped sinus has long been a taxonomic challenge, repeatedly discussed (e.g. Ryland 1974; Soule & Soule 1975). Examination of the types of W. subtorquata and W. cucullata has made possible to specify the diagnostic value of several morphological and morphometric characters, to better characterize these species and to correct previous records (Taylor & Gordon 2002; Chimenz Gusso et al. 2004; Vieira et al. 2014). Genetic studies have shown that cryptic diversity is remarkably high in Watersipora (Ryland et al. 2009; Mackie et al. 2006, 2012), obviously in relation with human-mediated dispersal on a wide geographic scale. Particularly in this genus, molecular approaches must be conducted jointly with precise examination of morphological characters. The Lebanese specimens of Watersipora were ascribed to W. subtorquata based on orificial criteria (Chimenz Gusso et al. 2004; Vieira et al. 2014)—a rounded sinus (more quadrangular in W. cucullata), small condyles and a mean width/length ratio of the orifice up to 1.14 in the Lebanese material, i.e. a value close to that of the type specimen (1.18: data from Soule & Soule 1975). The validity of this attribution was confirmed by L.M. Vieira (pers. comm., 19 February 2014). The distribution of W. subtorquata in the Mediterranean based on published literature and examination of museum specimens (Soule & Soule 1975; Chimenz Gusso et al. 2004; Abdel Salam & Ramadan 2008; Vieira et al. 2014) includes the eastern basin (Turkey, Israel, Egypt), the Central Mediterranean (Malta), the Western Mediterranean (Tyrrhenian and Ligurian seas: Civitavecchia, Naples, Genoa, Savona), and the Strait of Gibraltar (Tangiers). The features of specimens from Marseille stored at the Station Marine d’Endoume attest that W. subtorquata was established in the commercial port before the 1970 s, presumably introduced by vessels with fouled hulls., Published as part of Harmelin, Jean-Georges, 2014, Alien bryozoans in the eastern Mediterranean Sea — new records from the coast of Lebanon, pp. 301-338 in Zootaxa 3893 (3) on pages 314-315, DOI: 10.11646/zootaxa.3893.3.1, http://zenodo.org/record/250297, {"references":["Taylor, P. D. & Gordon, D. P. (2002) Alcide d'Orbigny's work on Recent and fossil bryozoans. Comptes Rendus Palevol, 1, 533 - 547. http: // dx. doi. org / 10.1016 / s 1631 - 0683 (02) 00067 - 2","Chimenz Gusso, C., Boccia, P. & Giovannini, N. (2004) Importance of faunistic and taxonomical studies for a correct analysis of the zoogeography of Mediterranean Bryozoa. Biogeographia, 25, 93 - 108.","Abdel-Salam, K. M. & Ramadan, S. E. (2008) Fouling Bryozoa from some Alexandria harbours, Egypt. (II) Encrusting species. Mediterranean Marine Science, 9, 5 - 20. http: // dx. doi. org / 10.12681 / mms. 129","Vieira, L. M., Spencer Jones, M. E. & Taylor, P. D. (2014) The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species. Zootaxa, 3857 (2), 151 - 182. http: // dx. doi. org / 10.11646 / zootaxa. 3857.2.1","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 Memoirs, 97, 1 - 158.","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.","Ryland, J. S., De Blauwe, H., Lord, R. & Mackie, J. A. (2009) Recent discoveries of alien Watersipora (Bryozoa) in Western Europe. Zootaxa, 2093, 43 - 59.","Norman, A. M. (1864) On undescribed British Hydrozoa, Actinozoa and Polyzoa. Annals and Magazine of Natural History, Series 3, 13, 82 - 90, pls. 9 - 11.","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, pp. i-viii, 55 - 120, pls. 69 - 124.","Waters, A. W. (1909) Reports on the marine biology of the Sudanese Red Sea. XII. The Bryozoa. Part I. - Cheilostomata. Journal of the Linnean Society, London, Zoology, 31 (205), 123 - 181. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1909. tb 00458. x","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.","Soule, D. F. & Soule, J. D. (1975) Species groups in Watersiporidae. In: Pouyet, S. (Ed.), Bryozoa 1974. Proceedings of the Third Conference, International Bryozoology Association. Documents des Laboratoires de Geologie de la Faculte des Sciences de Lyon, H. S. 3, pp. 299 - 309.","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. http: // dx. doi. org / 10.1007 / s 00227 - 005 - 0196 - x","Mackie, J. A., Darling, J. A. & Geller, J. B. (2012) Ecology of cryptic invasions: latitudinal segregation among Watersipora (Bryozoa) species. Scientific Reports, 2, 871. http: // dx. doi. org / 10.1038 / srep 00871"]}

Harmelin, Jean-Georges (2014): Alien bryozoans in the eastern Mediterranean Sea—new records from the coast of Lebanon. Zootaxa 3893 (3): 301-338, DOI: http://dx.doi.org/10.11646/zootaxa.3893.3.1

Watersipora subatra (Ortmann, 1890) n. stat. (Figs 39–53, 66, 69; Table 3) Schizoporella aterrima var. subatra Ortmann, 1890: 49 [Japan]. ? Watersipora cucullata var. watersi Mawatari, 1952: 12 (part), fig. 1G [Japan]. Dakaria subovoidea: Harmer 1957: p. 1022 (part), pl. 49, figs 11–12, 14 [Indonesia]. ? Cellepora subovoidea d’Orbigny, 1852: 402 [Red Sea; nomen dubium]. Watersipora subtorquata: Ryland 1974: 345, fig. 3C [Low Island]. Non Cellepora subtorquata d’Orbigny, 1852: 399 [Brazil]. “ Watersipora subovoidea ” fide Harmer: Soule & Soule 1975: 308 (part), pl. 3, fig. 6 [Australia]. “ Watersipora subovoidea ” fide Harmer: Winston & Heimberg, 1986: 15, figs 35–37 [Komodo]. Watersipora subtorquata: Gordon 1989: 40, pl. 20, B–H [New Zealand]. Watersipora subtorquata: Gordon & Mawatari, 1992: 30 [New Zealand]. Watersipora subtorquata: Ryland et al. 2009: 55, figs 3, 4A, C, E, F [Australia, Bay of Biscay and English Channel]. Watersipora subtorquata: Kelso & Wyse-Jackson 2012: 2010, fig. 1 [Ireland]. Watersipora subtorquata: Kuhlenkamp & Kind 2013: 3, figs 1B, C, 2A–F [North Sea: Helgoland]. Material examined. Holotype: MZS 002, Ortmann Collection, Sagami Bay, Japan, 50–100 fm (15–30 m), 1882. Other material: NHMUK 1922.10.8.3, dry slide, Schizoporella cucullata, sea level, Misaki, Japan, No. 1, Insole coll. NHMUK 1929.12.31.1, dry slide, Watersipora atrofusca, Australia, MacGillivray coll. NHMUK 2005.5.26.19, dry, Watersipora subtorquata, scraped from hull of Seasprite, Leigh Wharf / Marina, North Island, New Zealand, 6 October 2004, K. Tilbrook coll. NHMUK 2007.12.14.9, balsam slide, Watersipora subtorquata, J.S. Ryland det., Low Islands, Great Barrier Reef, July 1974, J.S. Ryland coll. NHMUK 2007.12.14.10, balsam slide, Watersipora subtorquata, J.S. Ryland det., Low Islands, Great Barrier Reef, July 1974, J.S. Ryland coll. NHMUK 2007.12.14.12, balsam slide, Watersipora subtorquata, J.S. Ryland don., on shell, Bay of Arcachon, August 2003, Hans de Blauwe coll. NHMUK 2007.12.14.14, balsam slide, Watersipora subtorquata, J.S. Ryland don., St-Jacut-sur-la-Mer, April 2005, Hans de Blauwe coll. NHMUK 2007.12.14.15, balsam slide, Watersipora subtorquata, J.S. Ryland don., St-Jacut-sur-la-Mer, April 2005, Hans de Blauwe coll. NHMUK 2007.12.14.16, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007, R. Lord coll. NHMUK 2007.12.14.17, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007, R. Lord coll. NHMUK 2007.12.14.18, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007, R. Lord coll. NHMUK 2010.12.1.2, dry, Watersipora sp., Point Loma, Marina, San Diego, California, USA, September 2010. P.D. Taylor & B. Okamura coll. NHMUK 2010.6.30.7, dry and dry slide, Long Beach, Los Angeles, USA. NHMUK 2012.6.30.6, dry slide [part of NHMUK 2005.5.26.19], Watersipora subtorquata, scraped from hull of Seasprite, hauled out Leigh Wharf / Marina, North Island, New Zealand, 6 October 2004, K. Tilbrook coll. Description. Colonies encrusting, multiserial, uni- to multilamellar; sometimes erect, foliaceous and bilamellar; colour in life variable, from orange to brownish-purple or greyish to black. Zooids subrectangular to hexagonal, about twice as long as wide, separated by slightly raised lateral walls. Frontal shield slightly convex, with numerous round pseudopores about 25 µm diameter; two large latero-oral intrazooidal septula near lateral zooidal margin, proximolateral to orifice, each with 3–8 (often 5) small pores. Frontal shield obscured by translucent cuticle. Orifice large, subcircular to oval, slightly wider than long, with broad, well-defined proximal, U-shaped sinus; orificial rim thin, sometimes slightly raised; narrow bar-shaped condyles, sometimes inconspicuous. Operculum with broad, biconcave proximal band, without lucidae. Avicularia absent. Ooecia absent. Remarks. Ortmann (1890) described and figured Schizoporella aterrima from Japan, including the new variety subatra without drawings or description. Ryland et al. (2009) noted that Schizoporella aterrima Ortmann, 1890 is unrecognizable and that the description given by Mawatari (1952) for Watersipora cucullata var. watersi, another species reported from Japan, may include at least two species. One of us (MSJ) has examined Ortmann’s specimens deposited at the Musée zoologique de la Ville de Strasbourg, concluding that they represent two species, Watersipora aterrima (MZS 003; Figs 35–38; Table 3) and Watersipora subatra (MZS 002; Fig. 39–42; Table 3). Watersipora aterrima is distinguished from W. subatra by having a smaller orifice, narrower sinus and smaller frontal pseudopores. Mawatari (1952) introduced variety watersi for specimens with distinctive opercula. His notes on this new variety are based on specimens from Japan, and two others recorded by Waters (1909) from Naples and by Hastings (1930) from Cape Verde. Unfortunately, Mawatari’s specimens from Japan assigned to W. cucullata var. watersi could not be located, but part of his description and figures resembles W. subatra and may refer to this species. We have assigned the Cape Verde and Naples specimens to a different species, Watersipora souleoroum n. sp. (see below). Comparisons between specimens from New Zealand (Figs 43–47), California (Fig 48–53) and Britain (some slides at NHMUK; see Ryland et al. 2009, fig. 3, as W. subovoidea) with the type specimens of W. cucullata, W. subatra and W. subtorquata, indicate that W. subatra was previously misidentified as W. subtorquata by some authors (e.g. Ryland 1974; Gordon 1989; Gordon & Mawatari 1992; Mackie et al. 2006, 2012; Geller et al. 2008; Ryland et al. 2009; Láruson et al. 2012; Cockrell & Sorte 2013; Needles & Wendt 2013; Sorte & White 2013; Davis & Marshall 2014). Since Ryland et al. (2009) found the COI haplotype in colonies from Guernsey and Brittany to be identical to the commonest haplotype identified in other areas, and the morphology of the genetically identified specimens to be the same (as W. subtorquata; Mackie et al. 2006), we believe that W. subatra is the commonest putatively invasive species of Watersipora in Britain, Australia, New Zealand and California, whereas W. subtorquata is the predominant species in subtropical and tropical waters of the Atlantic and Mediterranean. Colonies from California (Figs 48–53) often have a well-demarcated, U-shaped sinus and a smooth suborificial region (without pseudopores), as has been figured in specimens from Brittany (Ryland et al. 2009, fig. 3A, D, as W. subtorquata). All specimens assigned to W. subatra have an orifice with a U-shaped sinus and narrow, bar-shaped condyles, opercula with a broad, biconcave band proximally, frontal shields with pseudopores 18–30 µm in diameter, and two latero-oral intrazooidal septula. Soule and Soule (1975) indicated that Harmer (1957) included at least two species under Dakaria subovoidea, which may be a mixture of W. subtorquata and W. aterrima. According to the list of specimens published by Harmer (1957), however, his description and remarks included at least four species: W. subtorquata, W. cucullata, W. atrofusca and W. subatra (specimens figured from Siboga Stns 181 and 184, see Harmer 1957, pl. 49, figs 11–12, 14). Watersipora subatra is distinguished from W. subtorquata by the shapes of the orifice and condyles, and the presence of latero-oral intrazooidal septula (Figs 47, 52, 53). Latero-oral intrazooidal septula are also found in W. cucullata (Figs 8, 29, 34), W. aterrima (Fig. 37) and W. mawatarii n. sp. (Fig. 58; see below), but these species have different shapes of orifice and condyles. Despite the wide distribution of W. subatra in Pacific waters (see Ryland et al. 2009; Mackie et al. 2012; as W. subtorquata), its occurrence in some localities in the northwest Pacific, as well as the occurrence of W. subtorquata, are unconfirmed and require morphological and molecular investigation. Distribution. NE Atlantic (Ireland and British Isles), Indo-West Pacific (Indonesia) and Pacific (New Zealand, Australia and California).

Watersipora souleorum n. sp. (Figs 17, 59–64, 68, 71; Table 4) ? Cellepora subovoidea d’Orbigny, 1852: 402 [Red Sea; nomen dubium]. Lepralia cucullata: Thornely 1905: 120 [Sri Lanka]. Non Lepralia cucullata Busk, 1854: 81, pl. 96, figs 4–5 [Aegean Sea]. Schizoporella cucullata var. labiosa Calvet in Jullien & Calvet, 1903: 141, pl. 16, fig. 7a–c [Azores]. Lepralia? cucullata: Waters 1909: 150 (part), pl. 15, figs 2–5 [Naples and? Cape Verde]. Watersipora cucullata: Hastings 1930: 729 (part), pl. 15, fig. 104 [Cape Verde]. Watersipora cucullata var. watersi Mawatari, 1952: 12 (part) [Cape Verde]. “ Watersipora subovoidea ” fide Harmer: Soule & Soule 1975: 308 (part), pl. 3, fig. 5 [Sri Lanka]. Material examined. Holotype: NHMUK 2014.07.29.1, dry slide, Porto Grande, St. Vicent, Cape Verde Islands, on coral, 10 fm [18.3 m], Vallentin coll. Paratype: NHMUK 1935.3.6.367, same data as for holotype. Other material: MOM 420379, Schizoporella cucullata var. labiosa, L. Calvet 1903, l’Hirondelle, Stn 236, Azores, 20 August 1888. NHMUK 1872.2.3.147, dry, Watersipora cucullata var. atrofusca, west coast of Spain, S. Kent coll., on shells. NHMUK 1873.7.21.7, dry and dry slide, Watersipora cucullata, Ceylon [Sri Lanka], E.W.H. Holdsworth. NHMUK 1882.10.18.3.4–41pt., Watersipora sp., Seychelles Island, on coral, 12 fm [22 m], Voyage of H.M.S. Alert, pres. Lords of the Admiralty. NHMUK 1882.10.18.3.100–104., Schizoporella sp., Seychelles Island, coral, 12 fm [22 m], Voyage of H.M.S. Alert, pres. Lords of the Admiralty. NHMUK 1899.5.1.976 [3 slides], Watersipora atrofusca, Trincomalee, Sri Lanka, T. Hincks coll. NHMUK 1899.7.1.1396, dry slide, Lepralia cucullata, Persian Gulf, on pearl oyster, G. Busk coll. NHMUK 1899.7.1.1893, dry slide [part in balam slide], Lepralia cucullata, Cape Verde Island, Mrs Gatty, G. Busk coll. NHMUK 1890.1.31.13, dry, Watersipora subovoidea, Madras [Chennai], India. NHMUK 1912.12.21.1019, Watersipora cucullata, Naples, Italy, A.M. Norman. NHMUK 1921.5.24.18C, dry, Watersipora subovoidea, Cape Verde, Voyage of the Herald, Lords of the Admiralty. NHMUK 1936.12.30.35, Lepralia cucullata, Ceylon [Sri Lanka], L. R. Thornely. NHMUK 1936.12.30.116, Lepralia cucullata, India, No. 116, 1906, Yellow 2247, L. R. Thornely. NHMUK 1969.1.16.3, dry, Watersipora subvoidea, Point de Fomone, M’Bour, Senegal, 10 m, M-Marchad II. NHMUK 1973.1.10.2, dry slide, Watersipora subvoidea, No. 32, Point de Fomone, M’Bour, Senegal, 10 m, 13 April 1954, M-Marchad II. NHMUK 1981.4.1.3pt, dry and balsam slide, Watersipora subovoidea, Ceylon [Sri Lanka], L. R. Thornely, Rep. Pearl Oyster Fish., 26, p120. NHMUK 2012.6.30.1 [part of NHMUK 1912.12.21.1019], Watersipora cucullata, Naples, Italy, A.M. Norman. Description. Colonies encrusting, multiserial, uni- to multilamellar, sometimes erect, foliaceous and bilamellar; colour of dead colonies black. Zooids subrectangular, about twice as long as wide, separated by slightly raised lateral walls. Frontal shield slightly convex, with numerous (25–32 µm diameter), round pseudopores; latero-oral intrazooidal septula absent. Frontal shield obscured by dark cuticle. Orifice large, oval, with welldefined, broad, deeply U-shaped proximal sinus; slightly raised orificial rim sometimes present; bar-shaped condyles. Operculum with a narrow, biconcave dark central band, with lucidae. Avicularia absent. Ovicells absent. Etymology. Named for John D. Soule (1920–2001) and Dorothy F. Soule (1923–2005), in recognition of their contributions to bryozoan taxonomy. Remarks. The name labiosa was first introduced by Hincks (1886) —as Schizoporella atrofusca f. labiosa —but was synonymised under W. subovoidea by Ryland et al. (2009) (= W. subtorquata). Calvet (in Jullien & Calvet 1903) also introduced the name labiosa as Schizoporella cucullata var. labiosa —this name has never been considered valid since the original description and it was synonymised with W. cucullata by Hastings (1927) —for material from the Azores. Calvet’s specimens (MOM 420379) are distinct from W. subtorquata, however, in having a deep sinus and an operculum with a biconcave dark central band. Hastings (1930, pl. 15, fig. 104, as W. cucullata) included an excellent drawing based on specimens from Cape Verde that corresponds with Calvet’s variety. Waters (1909) used the name L. cucullata for species from Naples, Cape Verde and Suez, but the descriptions encompass at least two species. His specimens from Naples (Waters 1909, figs 2–5) have a wider sinus and distinctive operculum when compared to those from Suez (= W. subtorquata); specimens from Naples resemble those reported by Calvet (in Jullien & Calvet 1903) and Hastings (1930) from the Azores and Cape Verde, respectively. Mawatari (1952) erected var. watersi for material with this distinctive type of operculum, but the name is preoccupied by the fossil Microporella watersi de Stefani, 1884, based on Lepralia cucullata of Manzoni (1875) which, to judge by Manzoni’s figure (1875, pl. 4, fig. 47), is likely a species of Watersipora (see Waters 1909). We introduce the name Watersipora souleorum n. sp. for these specimens. Watersipora souleorum n. sp. resembles W. subtorquata in lacking latero-oral intrazooidal septula. The new species differs, however, in having a larger orifice than W. subtorquata, a deep U-shaped sinus, bar-shaped condyles, and an operculum with a narrow dark central band. Distribution. Atlantic (Azores, Cape Verde and Senegal), Mediterranean (Naples) and Indian Ocean (Sri Lanka and India).

Watersipora subtorquata (d’Orbigny, 1852) (Figs 1–5, 12–16, 18–24, 67, 70; Table 1) Escharina torquata: d’Orbigny, 1842: pl. 4, fig. 3; 1847: 11 [Brazil]. Cellepora subtorquata d’Orbigny, 1852: 399 [Brazil].?Non Cellepora subovoidea d’Orbigny, 1852: 402 [Red Sea; nomen dubium]. Schizoporella atrofusca: Hincks 1886: 269 (part; f. labiosa), pl. 10, fig. 5 (non fig. 4) [Arabian Sea]. Non Schizoporella atrofusca Busk, 1856: 178 [Mexico]. Dakaria chevreuxi Jullien in Jullien & Calvet, 1903: 90, pl. 10, fig. 6 [Senegal]. Lepralia ? cucullata: Waters 1909: 150 (part), pl. 15, fig. 1 [Suez]. Non Lepralia cucullata Busk, 1854: 81, pl. 96, figs 4–5 [Aegean Sea]. Watersipora cucullata: Hastings 1930: 729 (part), pl. 15, figs 102 [Suez]. Watersipora cucullata: Marcus 1937: 118, pl. 24, fig. 63A, B [Brazil]. Watersipora cucullata: Marcus 1938: 46 [Brazil]. Watersipora cucullata: Osburn 1952: 472 (part), pl. 56, fig. 4 [Colombia]. Dakaria subovoidea: Harmer 1957: 1022 (part). Watersipora edmondsoni Soule & Soule, 1968: 215, pl. 2, fig. 3 [Hawaii]. Watersipora subovoidea: Ryland 1974: 345, fig. 3A [SE Australia]. Watersipora subtorquata: Soule & Soule 1975: 302, pl. 3, fig. 3 [Brazil]; 304, pl. 2, fig. 3 [Virgin Islands]; pl. 2, fig. 5 [Bermuda]. Watersipora subovoidea fide Harmer: Soule & Soule 1975: 302 (part), pl. 3, fig. 4 [Alexandria]. Watersipora subovoidea: Winston 1982: 139, fig. 66 [Florida]. Watersipora subtorquata: d’Hondt 1988: 199, figs. 6.1–2 [Israel]. Watersipora subtorquata: Seo 1999: 222 (?part), fig. 1 [Korea]. Watersipora subtorquata: Taylor & Gordon 2002: 4 (text), fig. 1A–C [Brazil]. Watersipora subtorquata: Florence et al. 2007: 39, fig. 14I, J [South Africa]. Watersipora subtorquata: Abdel Salam & Ramadan 2008: 9, fig. 3 [Alexandria]. Watersipora subovoidea: Ryland et al. 2009: 54, figs. 4C, D, G, H [Australia and Italy]. Watersipora subovoidea: Ramalho et al. 2011: 772, fig. 3 [Brazil]. Material examined. Holotype: MNHN, d’Orbigny Collection 13637, Rio de Janeiro, Brazil. Other material: MZUSP 0257, Watersipora subtorquata, Araçá, São Sebastião, São Paulo, Brazil, 7 July 2009. NHMUK 1863.8.2.41, dry, Watersipora cucullata, Alexandria, Egypt, Station 34a, Eastern Harbour, O’Donoghue coll. NHMUK 1879.4.25.23, dry slide, Schizoporella atrofusca, Naples, Italy, A.W. Waters coll. NHMUK 1884.4.21.2, dry, Watersipora cucullata, South Chinese Seas. NHMUK 1885.1.26.6–7, dry, Watersipora subovoidea, Red Sea, A. Carpenter Esq. NHMUK 1888.11.14.110, dry slide, Schizoporella sp., Port Phillip Heads, Australia, January 1887, K. Bracebridge Wilson coll. NHMUK 1888.11.14.111, dry slide, Schizoporella cucullata, Port Phillip Heads, Australia, January 1887, J. Bracebride Wilson coll. NHMUK 1888.11.14.358, dry slide, Lepralia cucullata, Port Phillip Heads, Australia, January 1887, J. Bracebride Wilson coll. NHMUK 1899.5.1.974 [one dry slide and one balsam slide], Schizoporella atrofusca form. labiosa, Type, Arabian Sea, T. Hincks coll. NHMUK 1899.7.1.139, dry slide, Lepralia cucullata, John Adams Bank, H.M.S. Herald, G. Busk coll. NHMUK 1899.7.1.5061, dry slide, Phylactella sp., Pensian Gulf?, G. Busk coll. NHMUK 1899.7.1.5214, dry slide, Lepralia cucullata, Tangiers, Morocco, 35 fm [62.18 m], HMS Shearwater, G. Busk coll. NHMUK 1913.6.4.11, dry slide, Leraplia cucullata, Muscat, Oman, 8–10 fm [14.6–18.3 m], Major S.G. Knox. NHMUK 1926.9.6.164, dry slide, Lepralia cucullata, Suez Canal, K 13, 4 November 1924, A. Hastings. NHMUK 1926.9.6.165, dry slide, Lepralia cucullata, Suez Canal, K 13, 4 November 1924, A. Hastings. NHMUK 1926.9.6.166, balsam slide, Lepralia cucullata, Suez Canal, K 13, 4 November 1924, A. Hastings. NHMUK 1929.8.31.1, balsam slide, Watersipora cucullata, Shema Creek, Malta, Mediterranean. NHMUK 1948.2.16.28, wet, Watersipora cucullata, Santos, Brazil, E. Marcus coll. NHMUK 1968.1.16.77, dry, Watersipora cucullata, St James, South Africa, 15 February 1933, F78, O’Donoghue coll. NHMUK 1963.8.2.41pt, dry, Watersipora subovoidea, J. Soule & D. Soule det., Alexandria, Egypt, Station 34a, Eastern Harbour, O’Donoghue coll. NHMUK 1973.1.10.3, dry, Accra, Ghana, No. 13B, 14 February 1949, Bassindale coll. NHMUK 1973.1.10.4pt, dry slide, Watersipora cucullata, Pram Pram 0.25 m, Gold coast [Ghana], 6 February 1950, Bassindale coll. NHMUK 1980.1.12.3, dry slide, Watersipora subovoidea, Townsville harbour, Queensland, Australia, D. Hall. NHMUK 1981.4.1.2, dry slide, Schizoporella cucullata, Hurghada, Red Sea, Yellow 935. NHMUK 1986.8.14.3, dry slide, Watersipora subovoidea, Hutchinson Island, Florida, 14 November 1974, JWD coll. NHMUK 1996.9.4.1, dry, Watersipora subtorquata,? Site 4, Punta Mochila, Venezuela, October 1971, J.S. Ryland. NHMUK 1996.9.4.2, dry, Watersipora subtorquata, Site 1, West of Punto Arrecifes, West of Caracas, Venezuela, 8 October 1971, J.S. Ryland. NHMUK 1997.3.12.7, dry, Watersipora subtorquata, Mochina Bay, nr. Naiguatá, coast of Caracas, Venezuela, Loc. 2a, West shore of Gay, 11 October 1971, J.S. Ryland. NHMUK 1997.3.12.3, dry, Watersipora subtorquata, Site 5, Playa Caribe, Juan Griego, Venezuela, 12 October 1971, J.S. Ryland. NHMUK 1998.8.5.2, dry, Watersipora subtorquata, Mughsay’, near Salalah, Southern Oman, 10 October 1983, J.D. Taylor coll. NHMUK 2012.6.30.3, dry slide A/B [part of NHMUK 1998.8.5.2], Watersipora subtorquata, Mughsay’, near Salalah, Southern Oman, 10 October 1983, J.D. Taylor coll. NHMUK 1999.3.30.11, dry slide, Watersipora sp., New Caledonia, Beach of Ile du Phare (Lighthouse), Noumea, A. Willey. NHMUK 2007.12.14.2, balsam slide, Watersipora subovoidea, J.S. Ryland det., Naples, Italy, 20 October 1960. NHMUK 2007.12.14.3, balsam slide, Watersipora subovoidea, J.S. Ryland det., Arrawarra, New South Wales, Australia, January 1972. NHMUK 2007.12.14.4, balsam slide, Watersipora subovoidea, J.S. Ryland det., Arrawarra, New South Wales, Australia, January 1972. NHMUK 2007.12.14.5, balsam slide, Watersipora subovoidea, J.S. Ryland det., Arrawarra, New South Wales, Australia, January 1972, J.S. Ryland coll. NHMUK 2007.12.14.6, balsam slide, Watersipora subovoidea, J.S. Ryland det., Genoa Harbour, Italy, 12 November 2007, M. Faimali coll. NHMUK 2007.12.14.7, balsam slide, Watersipora subovoidea, J.S. Ryland det., Genoa Harbour, Italy, 12 November 2007, M. Faimali coll. NHMUK 2007.12.14.8, balsam slide, Watersipora subovoidea, J.S. Ryland det., Genoa Harbour, Italy, 12 November 2007, M. Faimali coll. NHMUK 2012.6.30.15, dry, Watersipora subovoidea, Shaab Baraja, Sudan, Red Sea, Large lump covered in soft coral, 23m, 15 September 1978, Dumont coll. NHMUK 2012.6.30.16, dry, Watersipora subovoidea, Shaab Baraja, Sudan, Red Sea, 25º52’N, 37º24’W, coral rock, 20 m, 22 September 1978, Dumont coll. NHMUK 1926.9.6.167, balsam slide, Lepralia cucullata, Suez Canal, K 2, 17 October 1924, A. Hastings. NHMUK 2012.6.30.2, dry slide [part of NHMUK 1973.1.10.3], Watersipora subovoidea, Accra, Ghana, No. 13B, 14 February 1949, Bassindale coll. NHMUK 2012.6.30.4, dry slide [part of 1970.6.1.23], Watersipora subovoidea, Mikhmoret, Israel, 16 October 1967, G. Eitan. Description. Colony encrusting, multiserial, primarily unilamellar on flat substrata, becoming multilamellar on irregular substrata, sometimes erect, foliaceous and bilamellar; orange (Figs 20–21) to brownish-purple or black in life, dried colonies becoming dark-orange to grey. Zooids subrectangular to hexagonal, about twice as long as wide, separated by slightly raised lateral walls. Frontal shield flat to slightly convex, sometimes raised more distally than proximally, with numerous round pseudopores about 25 µm in diameter; latero-oral and disto-oral pseudopores sometimes smaller than frontal pseudopores. Latero-oral intrazooidal septula absent. Frontal shield obscured by transparent cuticle in unbleached material. Orifice large, subcircular to oval, slightly wider than long, with well-defined proximal sinus; orificial rim often with raised subtriangular projections on each side of sinus; condyles triangular, projecting distomedially, tooth-like, sometimes inconspicuous and obscured by proximal raised projections. Operculum with broad, parallel-sided dark central band demarcated by sclerites; two lucidae proximally, adjacent to condyles. Avicularia absent. Ooecia absent; orange-red embryos brooded internally, filling most of coelom of maternal zooid. Polypides bright red-orange, lophophore with 20–22 tentacles. Remarks. We figure scanning electron micrographs of specimens from Mediterranean Israel (NHMUK 1970.6.1.23, Figs 12, 18), the Indian Ocean [Oman (NHMUK 1998.8.5.2, Fig. 13)] and the Atlantic [Ghana (NHMUK 1973.1.10.3, Figs 14–16), as well as from Brazil (MZUSP 0257; Fig. 19)] to show their similarity with the holotype of W. subtorquata (Figs 1–5). Ryland et al. (2009), followed by Ramalho et al. (2011), used the name W. subovoidea in error for this species. Watersipora subtorquata is characterized by a suborbicular orifice with a sinus demarcated by triangular condyles and an operculum with a parallel-sided dark band and two well-defined lucidae adjacent to the condyles. Ryland et al. (2009) suggested that Lepralia cucullata of Waters (1909) comprised two species. Based on the figured specimens we suggest that the Suez specimens belong to W. subtorquata and the Naples specimens (Waters 1909, pl. 15, figs 2–4) to an undescribed species. We also found a species from Naples in the NHMUK collection (NHMUK 1912.12.21.1019, Fig. 17) that differs from W. subtorquata in the operculum and in having a wider orifice compared with the Brazilian specimens; this specimen resembles Waters’s specimens from Naples, which are here reassigned to Watersipora souleorum n. sp. (see below). Mawatari (1952) erected the ‘variety’ watersi for specimens of Lepralia cucullata reported by Waters (1909), but this name is preoccupied by a fossil, Microporella watersi de Stefani, 1884, which is supposedly a species of Watersipora (see Waters 1909). Japanese specimens assigned to W. cucullata var. watersi by Mawatari (1952, fig. 1G) lack the triangular condyles and lucidae in the operculum and probably belong to Watersipora subatra (see below). Ryland et al. (2009) noted that the rim of the orifice in W. subtorquata (as W. subovoidea) may be elevated on either side of the sinus, a character easily recognizable in colonies on algae from Israel and Brazil (Figs 18–19). Gordon (1989) showed a range of orificial morphologies in New Zealand specimens identified as W. subovoidea, which all seem to represent one species (D.P. Gordon pers. comm. to LMV, 2013). He described two latero-oral intrazooidal septula in his specimens (Gordon 1989, p. 40), a character we also saw in specimens from New Zealand deposited at the NHMUK (Fig. 47). Thus, the New Zealand specimens are here reassigned to Watersipora subatra. Hincks (1886) and Calvet (in Jullien & Calvet 1903) both independently introduced ‘ labiosa ’: as Schizoporella atrofusca f. labiosa Hincks, 1886 (p. 269, pl. 10, fig. 5), from the Arabian Sea; and Schizoporella cucullata var. labiosa Calvet in Jullien & Calvet, 1903 (p. 141, pl. 16, fig 7), from Cape Verde. The specimens identified as form labiosa by Hincks (1886) are conspecific with other specimens examined from the Arabian Sea (Fig. 13) and belong to W. subtorquata. The specimens previously identified as var. labiosa from the Azores by Calvet (in Jullien & Calvet 1903) (MOM 420379) are distinct in having a deep sinus and an operculum with a narrow, biconcave dark central band; these specimens are badly preserved, but they may be conspecific with W. souleorum n. sp. (see below). Seo (1999, 2005) figured specimens of putative Watersipora subtorquata from Korea, with reddish-brown to black colonies (Seo 2005: pls 154–156). The size of the orifice in her specimens (Seo 2005: pl. 156B) resembles W. subtorquata but the condyles are sometimes minute and obscured by the proximal peristome. Material reported as W. subtorquata by Ryland et al. (2009) has shallow, bar-shaped condyles and an operculum with a proximal biconcave band, quite distinct from W. subtorquata and W. cucullata; these specimens are here identified as Watersipora subatra (see below). Examination of the holotype material of Watersipora edmondsoni Soule & Soule, 1968 (SBMNH 96310, AHF 149; Fig. 20–22) revealed it to be W. subtorquata. We believe, however, that the additional specimens examined by Soule & Soule (1968, 1975) probably include one or more additional species from the same locality [e.g. the specimen reported as W. edmondsoni (Soule & Soule 1975, pl. 1, fig. 7)]. Specimens recently reported as Watersipora subovoidea sensu Harmer by Dick et al. (2006) belong to an undescribed species that may be conspecific with W. edmondsoni sensu Soule & Soule, 1975 (non Soule & Soule, 1968). Distribution. Atlantic (Brazil, Caribbean, Virgin Islands, Florida, Cape Verde, Senegal, Ghana and South Africa), Mediterranean (Italy, Alexandria), Red Sea, Arabian Sea and Pacific (China Sea, Korea, Hawaii and Australia)., Published as part of Vieira, Leandro M., Jones, Mary Spencer & Taylor, Paul D., 2014, The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species, pp. 151-182 in Zootaxa 3857 (2) on pages 155-162, DOI: 10.11646/zootaxa.3857.2.1, http://zenodo.org/record/4929981, {"references":["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. http: // dx. doi. org / 10.1080 / 00222938609460142","Busk, G. (1856) Zoophytology. Quarterly Journal of Microscopical Science, 4, 176 - 179, pls 7 - 8.","Jullien, J. & Calvet, L. (1903) Bryozoaires provenant des campagnes de l'Hirondelle (1886 - 1888). Resultats des campagnes scientifiques accomplies sur son yacht par Albert 1 er, Prince Souverain de Monaco, 1903, 23, 1 - 188.","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. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1909. tb 00458. x","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.","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] http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1929. tb 01453. x","Marcus, E. (1937) Bryozoarios Marinhos Brasileiros, 1. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 1, 5 - 224.","Marcus, E. (1938) Bryozoarios Marinhos Brasileiros, 2. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 2, 1 - 196.","Osburn, R. C. (1952) Bryozoa of the Pacific coast of America, part 2, Cheilostomata - Ascophora. Allan Hancock Pacific Expeditions, 14, 271 - 611. http: // dx. doi. org / 10.5962 / bhl. title. 6542","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga-Expeditie, 28 d, xv, 641 - 1147, pls. 42 - 74.","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.","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.","Soule, D. F. & Soule, J. D. (1975) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Winston, J. E. (1982) Marine bryozoans (Ectoprocta) of the Indian River area, Florida. Bulletin of the American Museum of Natural History, 173, 99 - 176.","Seo, J. E. (1999) Taxonomic review of Korean Watersipora (Bryozoa, Gymnolaemata, Cheilostomata). Korean Journal of Systematic Zoology, 15, 221 - 229.","Taylor, P. D. & Gordon, D. P. (2002) Alcide d'Orbigny's work on Recent and fossil bryozoans. Compte Rendu Palevol, 1, 533 - 547. http: // dx. doi. org / 10.1016 / S 1631 - 0683 (02) 00067 - 2","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.","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.","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. http: // dx. doi. org / 10.1080 / 00222933.2010.535917","Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14 - 17.","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.","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.","Canu, F. & Bassler, R. S. (1930) The Bryozoa of the Galapagos Islands. Proceedings of the United States National Museum, 76, 1 - 78.","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."]}

Watersipora mawatarii n. sp. (Figs 54–58; Table 4) ? Dakaria typica Okada & Mawatari, 1937: 438, pl. 11, fig. 6; text-fig. 2 [Japan]. ? Dakaria subovoidea: Kubota & Mawatari, 1985: 203, fig. 4 [Japan]. Material examined. Holotype: NHMUK 2012.6.30.8, Oshoro Marine Station, near Otaru, Hokkaido, Japan, intertidal, 16 April 1996, M.J.Weedon coll. Paratypes: NHMUK 2012.6.30.9–11, same data as holotype; NHMUK 2012.6.30.12–13, 17, Otaru chiko, Japan, on dead barnacles near shore, 12 December 1996, T. Kato coll. NHMUK 2012.6.30.18, dry, Amakusa Marine Biological Laboratory, Kyushu, Japan, 40–50m, dredged, 22 October 1996, T. Kato coll. Etymology. Named for Japanese bryozoologist Prof. Shunsuke F. Mawatari. Description. Colonies encrusting, multiserial, uni- to multilamellar; sometimes erect, foliaceous and bilamellar; colour greyish to black in dead colonies. Zooids elongate-elliptical to rectangular, widest below the orificial area, about twice as long as wide, separated by slightly raised lateral walls; zooids arranged in quincuncial series. Frontal shield thick, granulated, slightly convex, with numerous small (10–15 µm diameter), round pseudopores covering entire surface except for suborificial region; two latero-oral intrazooidal septula, near lateral zooidal margin, proximolateral to orifice, each with 3–6 small pores. Frontal shield obscured by opaque, dark cuticle. Orifice large, transversely elliptical, usually conspicuoualy wider than long, with well-defined proximal broad sinus; orificial rim often thick and raised, but some zooids with thin, slightly raised rim; narrow bar-shaped condyles occupying entire proximal edge of orifice, sometimes projecting medially as triangular projection. Operculum black, mushroom-shaped; lucidae present. Avicularia absent. Ovicells absent. Remarks. The overall zooidal morphology of specimens here figured and described resemble some Japanese specimens—viz. Dakaria typica Okada & Mawatari, 1937 (= Watersipora typica) from Miyagi, and Dakaria subovoidea from Hokkaido (Kubota & Mawatari 1985)—but they are distinguished by having fewer pseudopores in the frontal shield than Watersipora mawatarii n. sp. The thinner circular area in the distal half of the operculum, characteristic of W. typica, is also absent in W. mawatarii n. sp. Watersipora mawatarii n. sp. resembles W. edmondsoni sensu Soule & Soule (1975) from Hawaii (the holotype of Watersipora edmondsoni belongs to Watersipora subtorquata, see above), characterized by its long zooids (about 0.80–1.20 mm long and 0.40–0.60 mm wide) and dark-brown mushroom-shaped opercular pigmentation, with a curved lower portion that fits into the sinus area. Soule & Soule (1975) also reported frontal shields with very small pseudopores like those of W. mawatarii n. sp. The shape of the zooids (widest below the orificial area) and the absence of pseudopores in the suborificial region were also described for some specimens from different Indo-West Pacific sites—Bali (Winston & Heimberg 1986; as Watersipora edmondsoni), Vanuatu (Tilbrook et al. 2001; as Watersipora subovoidea sensu Harmer) and Hawaii (Dick et al. 2006; as Watersipora subovoidea sensu Harmer)—which we believe belong to an undescribed species; these specimens differ from W. mawatarii n. sp. in having a narrower sinus and smaller orifice: 0.189–230 mm long and 0.238 –0.270 mm wide in W. mawatarii n. sp. versus 0.162 –0.216 mm long and 0.198 –0.234 mm wide in species from Bali (Winston & Heimberg 1986), and 0.15–0.18 mm long and 0.20–0.25 mm wide in specimens from Hawaii (Dick et al. 2006). Watersipora mawatarii n. sp. resembles W. subatra in having a U-shaped sinus, bar-shaped condyles, and two latero-oral intrazooidal septula. The two species differ, however, in the size of the pseudopores (smaller in W. mawatarii n. sp.) and condyles (more conspicuous in W. mawatarii n. sp.) and in the absence of pseudopores in the suborificial region (characteristic of W. mawatarii n. sp.). Distribution. Japan.

Watersipora mawatarii n. sp. (Figs 54–58; Table 4) ? Dakaria typica Okada & Mawatari, 1937: 438, pl. 11, fig. 6; text-fig. 2 [Japan]. ? Dakaria subovoidea: Kubota & Mawatari, 1985: 203, fig. 4 [Japan]. Material examined. Holotype: NHMUK 2012.6.30.8, Oshoro Marine Station, near Otaru, Hokkaido, Japan, intertidal, 16 April 1996, M.J.Weedon coll. Paratypes: NHMUK 2012.6.30.9–11, same data as holotype; NHMUK 2012.6.30.12–13, 17, Otaru chiko, Japan, on dead barnacles near shore, 12 December 1996, T. Kato coll. NHMUK 2012.6.30.18, dry, Amakusa Marine Biological Laboratory, Kyushu, Japan, 40–50m, dredged, 22 October 1996, T. Kato coll. Etymology. Named for Japanese bryozoologist Prof. Shunsuke F. Mawatari. Description. Colonies encrusting, multiserial, uni- to multilamellar; sometimes erect, foliaceous and bilamellar; colour greyish to black in dead colonies. Zooids elongate-elliptical to rectangular, widest below the orificial area, about twice as long as wide, separated by slightly raised lateral walls; zooids arranged in quincuncial series. Frontal shield thick, granulated, slightly convex, with numerous small (10–15 µm diameter), round pseudopores covering entire surface except for suborificial region; two latero-oral intrazooidal septula, near lateral zooidal margin, proximolateral to orifice, each with 3–6 small pores. Frontal shield obscured by opaque, dark cuticle. Orifice large, transversely elliptical, usually conspicuoualy wider than long, with well-defined proximal broad sinus; orificial rim often thick and raised, but some zooids with thin, slightly raised rim; narrow bar-shaped condyles occupying entire proximal edge of orifice, sometimes projecting medially as triangular projection. Operculum black, mushroom-shaped; lucidae present. Avicularia absent. Ovicells absent. Remarks. The overall zooidal morphology of specimens here figured and described resemble some Japanese specimens—viz. Dakaria typica Okada & Mawatari, 1937 (= Watersipora typica) from Miyagi, and Dakaria subovoidea from Hokkaido (Kubota & Mawatari 1985)—but they are distinguished by having fewer pseudopores in the frontal shield than Watersipora mawatarii n. sp. The thinner circular area in the distal half of the operculum, characteristic of W. typica, is also absent in W. mawatarii n. sp. Watersipora mawatarii n. sp. resembles W. edmondsoni sensu Soule & Soule (1975) from Hawaii (the holotype of Watersipora edmondsoni belongs to Watersipora subtorquata, see above), characterized by its long zooids (about 0.80–1.20 mm long and 0.40–0.60 mm wide) and dark-brown mushroom-shaped opercular pigmentation, with a curved lower portion that fits into the sinus area. Soule & Soule (1975) also reported frontal shields with very small pseudopores like those of W. mawatarii n. sp. The shape of the zooids (widest below the orificial area) and the absence of pseudopores in the suborificial region were also described for some specimens from different Indo-West Pacific sites—Bali (Winston & Heimberg 1986; as Watersipora edmondsoni), Vanuatu (Tilbrook et al. 2001; as Watersipora subovoidea sensu Harmer) and Hawaii (Dick et al. 2006; as Watersipora subovoidea sensu Harmer)—which we believe belong to an undescribed species; these specimens differ from W. mawatarii n. sp. in having a narrower sinus and smaller orifice: 0.189–230 mm long and 0.238 –0.270 mm wide in W. mawatarii n. sp. versus 0.162 –0.216 mm long and 0.198 –0.234 mm wide in species from Bali (Winston & Heimberg 1986), and 0.15–0.18 mm long and 0.20–0.25 mm wide in specimens from Hawaii (Dick et al. 2006). Watersipora mawatarii n. sp. resembles W. subatra in having a U-shaped sinus, bar-shaped condyles, and two latero-oral intrazooidal septula. The two species differ, however, in the size of the pseudopores (smaller in W. mawatarii n. sp.) and condyles (more conspicuous in W. mawatarii n. sp.) and in the absence of pseudopores in the suborificial region (characteristic of W. mawatarii n. sp.). Distribution. Japan., Published as part of Vieira, Leandro M., Jones, Mary Spencer & Taylor, Paul D., 2014, The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species, pp. 151-182 in Zootaxa 3857 (2) on pages 171-174, DOI: 10.11646/zootaxa.3857.2.1, http://zenodo.org/record/4929981, {"references":["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 Tohoku Imperial University, Biology, 11, 433 - 445.","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.","Soule, D. F. & Soule, J. D. (1975) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Winston, J. E. & Heimberg, B. F. (1986) Bryozoans from Bali, Lombok, and Komodo. American Museum Novitates, 2847, 1 - 49.","Tilbrook, K. J., Hayward, P. J. & Gordon, D. P. (2001) Cheilostomatous Bryozoa from Vanuatu. Zoological Journal of the Linnean Society, 131, 35 - 109. http: // dx. doi. org / 10.1006 / zjls. 2000.0249"]}

Vieira, Leandro M., Jones, Mary Spencer, Taylor, Paul D. (2014): The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species. Zootaxa 3857 (2): 151-182, DOI: http://dx.doi.org/10.11646/zootaxa.3857.2.1

Genus Watersipora Neviani, 1896 Watersipora Neviani, 1896: 120; Osburn 1952: 471; Gordon 1989: 40. Dakaria Jullien in Jullien & Calvet, 1903: 90; Harmer 1926: 1921 (part). Pachycleithonia Canu & Bassler, 1930: 25. Type species. Lepralia cucullata Busk, 1854, by original designation. Diagnosis. Colony encrusting, multiserial, uni- to multilamellar, or erect, foliaceous and bilamellar. Colony coloured reddish to black in life owing to pigmented epitheca. Autozooids subrectangular to hexagonal, separated by raised walls. Cryptocystidean frontal shield with numerous rounded pseudopores; latero-oral intrazooidal septula sometimes present proximolateral to orifice; intrazooidal septula sometimes present at proximal corners of frontal shield. Orifice subcircular to oval; poster sometimes with well-defined proximal sinus; condyles present. Operculum reddish-brown to black in colour, often with central band demarcated by sclerites; lucidae often present. Spines absent. Avicularia absent. Ooecia absent; embryos brooded internally in maternal zooid. Multiporous mural pore plates in distolateral and transverse distal walls. Ancestrula schizoporelloid, single, smaller than autozooids, often obscured in later astogeny. Remarks. Watersipora was introduced monotypically for “ Smittia cucullata Busk, 1854 ” (= Lepralia cucullata Busk, 1854) by Neviani (1896) who stated that his fossil specimens were morphologically distinct from Lepralia cucullata. As a consequence, Harmer (1957) suggested using Dakaria Jullien in Jullien & Calvet, 1903 (type species: Dakaria chevreuxi Jullien in Jullien & Calvet, 1903, now Watersipora subtorquata; see below) rather than Watersipora for Busk’s species. Despite Neviani’s misidentification of his fossil specimens as Smittia (Watersipora) cucullata (Neviani 1896, p. 120; Gordon 1989, p. 40), Lepralia cucullata sensu Busk, 1854 is best used as the type species of Watersipora for the purpose of stability (see ICZN 1999, Article 70.3). Pachycleithonia Canu & Bassler, 1930 was introduced monotypically for Pachycleithonia nigra Canu & Bassler, 1930, from the Galapagos. The genus has been characterized as having ovicells (Cook 1985), but the ovicellate specimens are distinct from Canu & Bassler’s P. nigra, and Tilbrook (2006) subsequently reassigned them to Nigropercula Tilbrook, 2006. Osburn (1952) synonymised Pachycleithonia with Watersipora, but included Pachycleithonia nigra under the name Watersipora cucullata. We have examined the type material of Pachycleithonia nigra (USNM 8495; Fig. 11) and additional material from the Galapagos (NHMUK 1975.5.12.1); this species resembles W. cucullata in colony shape and in having a thick-rimmed orifice and triangular condyles. However, whereas Pachycleithonia nigra has a sinusoid orifice with a convex proximal edge, W. cucullata has a straight or slightly concave proximal edge, as found in the majority of species assigned to Watersipora, and paired intrazooidal (frontal) septula, which are lacking in Pachycleithonia nigra. These two characters in Pachycleithonia nigra also occur, however, in other Watersipora species, e.g. Watersipora subtorquata, which lacks paired intrazooidal septula, and Watersipora arcuata Banta, 1969a, which has an orifice with a convex proximal edge. Following Soule (1961), we recognize the combination Watersipora nigra (Canu & Bassler, 1930). Two other genera have been assigned to the family Watersiporidae. Veleroa Osburn, 1952 is distinct in having numerous communication pores covering the surface of the lateral and distal walls (see Osburn 1952: pl. 57, fig. 7), and Uscia Banta, 1969b has dimorphic ovicellate zooids (see Banta 1969b, figs 2–4)., Published as part of Vieira, Leandro M., Jones, Mary Spencer & Taylor, Paul D., 2014, The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species, pp. 151-182 in Zootaxa 3857 (2) on page 154, DOI: 10.11646/zootaxa.3857.2.1, http://zenodo.org/record/4929981, {"references":["Neviani, A. (1896) Briozoi fossili della Farnesina e Monte Mario presso Roma. Palaeontographia italica, 1, 77 - 140.","Osburn, R. C. (1952) Bryozoa of the Pacific coast of America, part 2, Cheilostomata - Ascophora. Allan Hancock Pacific Expeditions, 14, 271 - 611. http: // dx. doi. org / 10.5962 / bhl. title. 6542","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.","Jullien, J. & Calvet, L. (1903) Bryozoaires provenant des campagnes de l'Hirondelle (1886 - 1888). Resultats des campagnes scientifiques accomplies sur son yacht par Albert 1 er, Prince Souverain de Monaco, 1903, 23, 1 - 188.","Canu, F. & Bassler, R. S. (1930) The Bryozoa of the Galapagos Islands. Proceedings of the United States National Museum, 76, 1 - 78.","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.","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga-Expeditie, 28 d, xv, 641 - 1147, pls. 42 - 74.","ICZN (1999) International Code of Zoological Nomenclature. 4 th Edition. International Trust for Zoological Nomenclature, London, xx + 365 pp.","Cook, P. L. (1985) Bryozoa from Ghana. Tervuren, Belgie Koninklijk Museum voor Midden-Afrika. Zoologische Weteschappen, 238, 1 - 315.","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.","Banta, W. C. (1969 a) Watersipora arcuata, a new species in the subovoidea-cucullata-nigra complex (Bryozoa, Cheilostomata). Bulletin of the Southern California Academy of Science, 68, 96 - 102.","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.","Banta, W. C. (1969 b) Uscia mexicana, new genus, new species, a watersiporid bryozoan with dimorphic autozooids. Bulletin of the Southern California Academy of Science, 68, 20 - 35."]}

Identity of Watersipora cucullata (Busk, 1854) Busk (1854) described Lepralia cucullata from the Aegean Sea and noted that, owing to the absence of ovicells, it was not possible to assign his specimens to Savigny’s species (Savigny 1817, pl. 8, fig. 6), which had been named Cellepora mangnevillana Lamouroux, 1816 by Audouin (1826). The species was characterized by a black colouration and a granular surface (Busk 1854), and Waters (1879) described frontal pseudopores in Busk’s specimens as well as his own colonies from Naples. Hincks (1886) used the name Schizoporella atrofusca Busk, 1856 for the specimens reported by Waters (1879) and included the form labiosa from the Arabian Sea, but gave no additional accounts of Busk’s types of L. cucullata or S. atrofusca. Waters (1909) and Hastings (1927, 1930) also included S. atrofusca under L. cucullata, but their description encompassed more than one species (see also Soule & Soule 1975). Harmer (1957) synonymised three species— Cellepora subtorquata, Lepralia cucullata and Schizoporella atrofusca —under Watersipora subovoidea (as Dakaria subovoidea), which contrasts with the statement of Hastings (1930), who suggested the existence of more than one species under the name Watersipora cucullata; nevertheless, Hastings did not introduce new names for specimens with distinct opercular shapes. Mawatari (1952), however, introduced the variety watersi for specimens reported by Waters (1909) and Hastings (1930), which may represent at least two distinct species (see below). Following examination of the type specimens of Busk’s L. cucullata —i.e. Watersipora cucullata, NHMUK 1854.11.15.189, lectotype (Figs 6–9), chosen by Soule & Soule (1975), plus NHMUK 2012.6.30.1, paralectotype (Figs 25–29)—and S. atrofusca —i.e. Watersipora atrofusca, NHMUK 1892.9.6.4 (Fig. 10)—we conclude that these should be considered as distinct species, as was suggested by Soule & Soule (1975). Orifice size in the two species differs, being larger in W. cucullata than in W. atrofusca. Two small latero-oral multiporous septula in the frontal shield (equivalent to the ‘intrazoidal septula’ of Banta 1970, p. 39), one on each side of the orificial sinus, are present in W. cucullata (Figures 8, 29) but are lacking in W. atrofusca. Watersipora subtorquata is distinguished from Busk’s W. cucullata by the shape of the orifice, the shape and size of the condyles, and the size of the frontal pseudopores. Ryland et al. (2009) considered the identity of Busk’s W. cucullata “clear”, following the descriptions of Hastings (1930), but Hastings mentioned a distinct variety among her specimens, as noted in her description and plates (Hastings 1930, p. 730, pls 102–104). Ryland et al. (2009, p. 53) attempted to solve nomenclatural problems concerning W. subovoidea and to stabilise d’Orbigny’s species name by designating Busk’s type of W. cucullata as the neotype of d’Orbigny’s W. subovoidea, thereby making Watersipora subovoidea the senior synonym of Watersipora cucullata. Unfortunately, the descriptions and figures of W. subovoidea in Ryland et al. (2009) are actually of W. subtorquata (see above). Furthermore, the selection of a neotype for W. subovoidea from a site (Aegean Sea) distant from the type locality (Egyptian Red Sea) is contentious. According to the ICZN (1999, Article 75.3.6) evidence should be provided that the neotype came as nearly as practicable from the original type locality, which was not provided in the Ryland et al. (2009) paper. Thus, Cellepora subovoidea d’Orbigny, 1852 is here considered a nomen dubium. Below we provide redescriptions of d’Orbigny’s species W. subtorquata and W. cucullata and reassign the species erroneously identified as Watersipora subtorquata by Ryland et al. (2009) to a third species previously described by Ortmann (1890) under the name Schizoporella aterrima var. subatra (= Watersipora subatra, see below). We also describe two new species: Watersipora mawatarii n. sp. from Japan, and Watersipora souleorum n. sp. for specimens from Cape Verde and Naples previously assigned to Watersipora cucullata., Published as part of Vieira, Leandro M., Jones, Mary Spencer & Taylor, Paul D., 2014, The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species, pp. 151-182 in Zootaxa 3857 (2) on pages 153-154, DOI: 10.11646/zootaxa.3857.2.1, http://zenodo.org/record/4929981, {"references":["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.","Savigny, J. C. (1817) Description de l'Egypte, ou receuil des observations et des recherches qui ont ete faites en Egypte pendant l'Expedition de ' Armee francaise ... Histoire naturelle. Vol. 2. Imprimerie Royale, Paris, 125 pls.","Lamouroux, J. V. F. (1816) Histoire des Polypiers coralligenes flexibles, vulgairement nommes Zoophytes. F. Poisson, Caen, 559 pp.","Audouin, J. V. (1826) Explication sommaire des planches de Polypes de l'Egypte et de la Syrie, publiees par Jules-Cesar Savigny. In: Panckoucke, C. L. F. (Org.), Description de l'Egypte ou receuil des observations et des recherches qui ont ete faites en Egyptes pendant l'Expedition de l'Armee francaise … Histoire naturelle. Tome 1 (4). Imprimerie Imperiale, Paris, pp. 225 - 244.","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. http: // dx. doi. org / 10.1080 / 00222937908682488","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. http: // dx. doi. org / 10.1080 / 00222938609460142","Busk, G. (1856) Zoophytology. Quarterly Journal of Microscopical Science, 4, 176 - 179, pls 7 - 8.","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. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1909. tb 00458. x","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. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1927. tb 00381. x","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] http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1929. tb 01453. x","Soule, D. F. & Soule, J. D. (1975) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga-Expeditie, 28 d, xv, 641 - 1147, pls. 42 - 74.","Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14 - 17.","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. http: // dx. doi. org / 10.1002 / jmor. 1051310104","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.","ICZN (1999) International Code of Zoological Nomenclature. 4 th Edition. International Trust for Zoological Nomenclature, London, xx + 365 pp.","Ortmann, A. (1890) Die Japanische Bryozoenfauna. Bericht uber die von Herrn Dr. L. Doderlein im Jahre 1880 - 81 gemachten Sammlungen. Archiv fur Naturgeschichte, 54, 1 - 74."]}

Watersipora subatra (Ortmann, 1890) n. stat. (Figs 39–53, 66, 69; Table 3) Schizoporella aterrima var. subatra Ortmann, 1890: 49 [Japan]. ? Watersipora cucullata var. watersi Mawatari, 1952: 12 (part), fig. 1G [Japan]. Dakaria subovoidea: Harmer 1957: p. 1022 (part), pl. 49, figs 11–12, 14 [Indonesia]. ? Cellepora subovoidea d’Orbigny, 1852: 402 [Red Sea; nomen dubium]. Watersipora subtorquata: Ryland 1974: 345, fig. 3C [Low Island]. Non Cellepora subtorquata d’Orbigny, 1852: 399 [Brazil]. “ Watersipora subovoidea ” fide Harmer: Soule & Soule 1975: 308 (part), pl. 3, fig. 6 [Australia]. “ Watersipora subovoidea ” fide Harmer: Winston & Heimberg, 1986: 15, figs 35–37 [Komodo]. Watersipora subtorquata: Gordon 1989: 40, pl. 20, B–H [New Zealand]. Watersipora subtorquata: Gordon & Mawatari, 1992: 30 [New Zealand]. Watersipora subtorquata: Ryland et al. 2009: 55, figs 3, 4A, C, E, F [Australia, Bay of Biscay and English Channel]. Watersipora subtorquata: Kelso & Wyse-Jackson 2012: 2010, fig. 1 [Ireland]. Watersipora subtorquata: Kuhlenkamp & Kind 2013: 3, figs 1B, C, 2A–F [North Sea: Helgoland]. Material examined. Holotype: MZS 002, Ortmann Collection, Sagami Bay, Japan, 50–100 fm (15–30 m), 1882. Other material: NHMUK 1922.10.8.3, dry slide, Schizoporella cucullata, sea level, Misaki, Japan, No. 1, Insole coll. NHMUK 1929.12.31.1, dry slide, Watersipora atrofusca, Australia, MacGillivray coll. NHMUK 2005.5.26.19, dry, Watersipora subtorquata, scraped from hull of Seasprite, Leigh Wharf / Marina, North Island, New Zealand, 6 October 2004, K. Tilbrook coll. NHMUK 2007.12.14.9, balsam slide, Watersipora subtorquata, J.S. Ryland det., Low Islands, Great Barrier Reef, July 1974, J.S. Ryland coll. NHMUK 2007.12.14.10, balsam slide, Watersipora subtorquata, J.S. Ryland det., Low Islands, Great Barrier Reef, July 1974, J.S. Ryland coll. NHMUK 2007.12.14.12, balsam slide, Watersipora subtorquata, J.S. Ryland don., on shell, Bay of Arcachon, August 2003, Hans de Blauwe coll. NHMUK 2007.12.14.14, balsam slide, Watersipora subtorquata, J.S. Ryland don., St-Jacut-sur-la-Mer, April 2005, Hans de Blauwe coll. NHMUK 2007.12.14.15, balsam slide, Watersipora subtorquata, J.S. Ryland don., St-Jacut-sur-la-Mer, April 2005, Hans de Blauwe coll. NHMUK 2007.12.14.16, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007, R. Lord coll. NHMUK 2007.12.14.17, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007, R. Lord coll. NHMUK 2007.12.14.18, balsam slide, Watersipora subtorquata, J.S. Ryland don., Guersey, 2007, R. Lord coll. NHMUK 2010.12.1.2, dry, Watersipora sp., Point Loma, Marina, San Diego, California, USA, September 2010. P.D. Taylor & B. Okamura coll. NHMUK 2010.6.30.7, dry and dry slide, Long Beach, Los Angeles, USA. NHMUK 2012.6.30.6, dry slide [part of NHMUK 2005.5.26.19], Watersipora subtorquata, scraped from hull of Seasprite, hauled out Leigh Wharf / Marina, North Island, New Zealand, 6 October 2004, K. Tilbrook coll. Description. Colonies encrusting, multiserial, uni- to multilamellar; sometimes erect, foliaceous and bilamellar; colour in life variable, from orange to brownish-purple or greyish to black. Zooids subrectangular to hexagonal, about twice as long as wide, separated by slightly raised lateral walls. Frontal shield slightly convex, with numerous round pseudopores about 25 µm diameter; two large latero-oral intrazooidal septula near lateral zooidal margin, proximolateral to orifice, each with 3–8 (often 5) small pores. Frontal shield obscured by translucent cuticle. Orifice large, subcircular to oval, slightly wider than long, with broad, well-defined proximal, U-shaped sinus; orificial rim thin, sometimes slightly raised; narrow bar-shaped condyles, sometimes inconspicuous. Operculum with broad, biconcave proximal band, without lucidae. Avicularia absent. Ooecia absent. Remarks. Ortmann (1890) described and figured Schizoporella aterrima from Japan, including the new variety subatra without drawings or description. Ryland et al. (2009) noted that Schizoporella aterrima Ortmann, 1890 is unrecognizable and that the description given by Mawatari (1952) for Watersipora cucullata var. watersi, another species reported from Japan, may include at least two species. One of us (MSJ) has examined Ortmann’s specimens deposited at the Musée zoologique de la Ville de Strasbourg, concluding that they represent two species, Watersipora aterrima (MZS 003; Figs 35–38; Table 3) and Watersipora subatra (MZS 002; Fig. 39–42; Table 3). Watersipora aterrima is distinguished from W. subatra by having a smaller orifice, narrower sinus and smaller frontal pseudopores. Mawatari (1952) introduced variety watersi for specimens with distinctive opercula. His notes on this new variety are based on specimens from Japan, and two others recorded by Waters (1909) from Naples and by Hastings (1930) from Cape Verde. Unfortunately, Mawatari’s specimens from Japan assigned to W. cucullata var. watersi could not be located, but part of his description and figures resembles W. subatra and may refer to this species. We have assigned the Cape Verde and Naples specimens to a different species, Watersipora souleoroum n. sp. (see below). Comparisons between specimens from New Zealand (Figs 43–47), California (Fig 48–53) and Britain (some slides at NHMUK; see Ryland et al. 2009, fig. 3, as W. subovoidea) with the type specimens of W. cucullata, W. subatra and W. subtorquata, indicate that W. subatra was previously misidentified as W. subtorquata by some authors (e.g. Ryland 1974; Gordon 1989; Gordon & Mawatari 1992; Mackie et al. 2006, 2012; Geller et al. 2008; Ryland et al. 2009; Láruson et al. 2012; Cockrell & Sorte 2013; Needles & Wendt 2013; Sorte & White 2013; Davis & Marshall 2014). Since Ryland et al. (2009) found the COI haplotype in colonies from Guernsey and Brittany to be identical to the commonest haplotype identified in other areas, and the morphology of the genetically identified specimens to be the same (as W. subtorquata; Mackie et al. 2006), we believe that W. subatra is the commonest putatively invasive species of Watersipora in Britain, Australia, New Zealand and California, whereas W. subtorquata is the predominant species in subtropical and tropical waters of the Atlantic and Mediterranean. Colonies from California (Figs 48–53) often have a well-demarcated, U-shaped sinus and a smooth suborificial region (without pseudopores), as has been figured in specimens from Brittany (Ryland et al. 2009, fig. 3A, D, as W. subtorquata). All specimens assigned to W. subatra have an orifice with a U-shaped sinus and narrow, bar-shaped condyles, opercula with a broad, biconcave band proximally, frontal shields with pseudopores 18–30 µm in diameter, and two latero-oral intrazooidal septula. Soule and Soule (1975) indicated that Harmer (1957) included at least two species under Dakaria subovoidea, which may be a mixture of W. subtorquata and W. aterrima. According to the list of specimens published by Harmer (1957), however, his description and remarks included at least four species: W. subtorquata, W. cucullata, W. atrofusca and W. subatra (specimens figured from Siboga Stns 181 and 184, see Harmer 1957, pl. 49, figs 11–12, 14). Watersipora subatra is distinguished from W. subtorquata by the shapes of the orifice and condyles, and the presence of latero-oral intrazooidal septula (Figs 47, 52, 53). Latero-oral intrazooidal septula are also found in W. cucullata (Figs 8, 29, 34), W. aterrima (Fig. 37) and W. mawatarii n. sp. (Fig. 58; see below), but these species have different shapes of orifice and condyles. Despite the wide distribution of W. subatra in Pacific waters (see Ryland et al. 2009; Mackie et al. 2012; as W. subtorquata), its occurrence in some localities in the northwest Pacific, as well as the occurrence of W. subtorquata, are unconfirmed and require morphological and molecular investigation. Distribution. NE Atlantic (Ireland and British Isles), Indo-West Pacific (Indonesia) and Pacific (New Zealand, Australia and California)., Published as part of Vieira, Leandro M., Jones, Mary Spencer & Taylor, Paul D., 2014, The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species, pp. 151-182 in Zootaxa 3857 (2) on pages 166-171, DOI: 10.11646/zootaxa.3857.2.1, http://zenodo.org/record/4929981, {"references":["Ortmann, A. (1890) Die Japanische Bryozoenfauna. Bericht uber die von Herrn Dr. L. Doderlein im Jahre 1880 - 81 gemachten Sammlungen. Archiv fur Naturgeschichte, 54, 1 - 74.","Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14 - 17.","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga-Expeditie, 28 d, xv, 641 - 1147, pls. 42 - 74.","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.","Soule, D. F. & Soule, J. D. (1975) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Winston, J. E. & Heimberg, B. F. (1986) Bryozoans from Bali, Lombok, and Komodo. American Museum Novitates, 2847, 1 - 49.","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.","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.","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, e 73.","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. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1909. tb 00458. x","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] http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1929. tb 01453. x","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. http: // dx. doi. org / 10.1007 / s 00227 - 005 - 0196 - x","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.","Laruson, 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. http: // dx. doi. org / 10.1007 / s 12686 - 012 - 9606 - 9","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. http: // dx. doi. org / 10.1016 / j. jembe. 2012.11.008","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. http: // dx. doi. org / 10.1007 / s 10530 - 012 - 0362 - 2","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. http: // dx. doi. org / 10.1098 / rspb. 2013.0572","Davis, K. & Marshall, D. J. (2014) Offspring size in a resident species affects community assembly. Journal of Animal Ecology, 83, 322 - 331. http: // dx. doi. org / 10.1111 / 1365 - 2656.12136","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. http: // dx. doi. org / 10.1038 / srep 00871"]}

Watersipora souleorum n. sp. (Figs 17, 59–64, 68, 71; Table 4) ? Cellepora subovoidea d’Orbigny, 1852: 402 [Red Sea; nomen dubium]. Lepralia cucullata: Thornely 1905: 120 [Sri Lanka]. Non Lepralia cucullata Busk, 1854: 81, pl. 96, figs 4–5 [Aegean Sea]. Schizoporella cucullata var. labiosa Calvet in Jullien & Calvet, 1903: 141, pl. 16, fig. 7a–c [Azores]. Lepralia? cucullata: Waters 1909: 150 (part), pl. 15, figs 2–5 [Naples and? Cape Verde]. Watersipora cucullata: Hastings 1930: 729 (part), pl. 15, fig. 104 [Cape Verde]. Watersipora cucullata var. watersi Mawatari, 1952: 12 (part) [Cape Verde]. “ Watersipora subovoidea ” fide Harmer: Soule & Soule 1975: 308 (part), pl. 3, fig. 5 [Sri Lanka]. Material examined. Holotype: NHMUK 2014.07.29.1, dry slide, Porto Grande, St. Vicent, Cape Verde Islands, on coral, 10 fm [18.3 m], Vallentin coll. Paratype: NHMUK 1935.3.6.367, same data as for holotype. Other material: MOM 420379, Schizoporella cucullata var. labiosa, L. Calvet 1903, l’Hirondelle, Stn 236, Azores, 20 August 1888. NHMUK 1872.2.3.147, dry, Watersipora cucullata var. atrofusca, west coast of Spain, S. Kent coll., on shells. NHMUK 1873.7.21.7, dry and dry slide, Watersipora cucullata, Ceylon [Sri Lanka], E.W.H. Holdsworth. NHMUK 1882.10.18.3.4–41pt., Watersipora sp., Seychelles Island, on coral, 12 fm [22 m], Voyage of H.M.S. Alert, pres. Lords of the Admiralty. NHMUK 1882.10.18.3.100–104., Schizoporella sp., Seychelles Island, coral, 12 fm [22 m], Voyage of H.M.S. Alert, pres. Lords of the Admiralty. NHMUK 1899.5.1.976 [3 slides], Watersipora atrofusca, Trincomalee, Sri Lanka, T. Hincks coll. NHMUK 1899.7.1.1396, dry slide, Lepralia cucullata, Persian Gulf, on pearl oyster, G. Busk coll. NHMUK 1899.7.1.1893, dry slide [part in balam slide], Lepralia cucullata, Cape Verde Island, Mrs Gatty, G. Busk coll. NHMUK 1890.1.31.13, dry, Watersipora subovoidea, Madras [Chennai], India. NHMUK 1912.12.21.1019, Watersipora cucullata, Naples, Italy, A.M. Norman. NHMUK 1921.5.24.18C, dry, Watersipora subovoidea, Cape Verde, Voyage of the Herald, Lords of the Admiralty. NHMUK 1936.12.30.35, Lepralia cucullata, Ceylon [Sri Lanka], L. R. Thornely. NHMUK 1936.12.30.116, Lepralia cucullata, India, No. 116, 1906, Yellow 2247, L. R. Thornely. NHMUK 1969.1.16.3, dry, Watersipora subvoidea, Point de Fomone, M’Bour, Senegal, 10 m, M-Marchad II. NHMUK 1973.1.10.2, dry slide, Watersipora subvoidea, No. 32, Point de Fomone, M’Bour, Senegal, 10 m, 13 April 1954, M-Marchad II. NHMUK 1981.4.1.3pt, dry and balsam slide, Watersipora subovoidea, Ceylon [Sri Lanka], L. R. Thornely, Rep. Pearl Oyster Fish., 26, p120. NHMUK 2012.6.30.1 [part of NHMUK 1912.12.21.1019], Watersipora cucullata, Naples, Italy, A.M. Norman. Description. Colonies encrusting, multiserial, uni- to multilamellar, sometimes erect, foliaceous and bilamellar; colour of dead colonies black. Zooids subrectangular, about twice as long as wide, separated by slightly raised lateral walls. Frontal shield slightly convex, with numerous (25–32 µm diameter), round pseudopores; latero-oral intrazooidal septula absent. Frontal shield obscured by dark cuticle. Orifice large, oval, with welldefined, broad, deeply U-shaped proximal sinus; slightly raised orificial rim sometimes present; bar-shaped condyles. Operculum with a narrow, biconcave dark central band, with lucidae. Avicularia absent. Ovicells absent. Etymology. Named for John D. Soule (1920–2001) and Dorothy F. Soule (1923–2005), in recognition of their contributions to bryozoan taxonomy. Remarks. The name labiosa was first introduced by Hincks (1886) —as Schizoporella atrofusca f. labiosa —but was synonymised under W. subovoidea by Ryland et al. (2009) (= W. subtorquata). Calvet (in Jullien & Calvet 1903) also introduced the name labiosa as Schizoporella cucullata var. labiosa —this name has never been considered valid since the original description and it was synonymised with W. cucullata by Hastings (1927) —for material from the Azores. Calvet’s specimens (MOM 420379) are distinct from W. subtorquata, however, in having a deep sinus and an operculum with a biconcave dark central band. Hastings (1930, pl. 15, fig. 104, as W. cucullata) included an excellent drawing based on specimens from Cape Verde that corresponds with Calvet’s variety. Waters (1909) used the name L. cucullata for species from Naples, Cape Verde and Suez, but the descriptions encompass at least two species. His specimens from Naples (Waters 1909, figs 2–5) have a wider sinus and distinctive operculum when compared to those from Suez (= W. subtorquata); specimens from Naples resemble those reported by Calvet (in Jullien & Calvet 1903) and Hastings (1930) from the Azores and Cape Verde, respectively. Mawatari (1952) erected var. watersi for material with this distinctive type of operculum, but the name is preoccupied by the fossil Microporella watersi de Stefani, 1884, based on Lepralia cucullata of Manzoni (1875) which, to judge by Manzoni’s figure (1875, pl. 4, fig. 47), is likely a species of Watersipora (see Waters 1909). We introduce the name Watersipora souleorum n. sp. for these specimens. Watersipora souleorum n. sp. resembles W. subtorquata in lacking latero-oral intrazooidal septula. The new species differs, however, in having a larger orifice than W. subtorquata, a deep U-shaped sinus, bar-shaped condyles, and an operculum with a narrow dark central band. Distribution. Atlantic (Azores, Cape Verde and Senegal), Mediterranean (Naples) and Indian Ocean (Sri Lanka and India)., Published as part of Vieira, Leandro M., Jones, Mary Spencer & Taylor, Paul D., 2014, The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species, pp. 151-182 in Zootaxa 3857 (2) on pages 174-177, DOI: 10.11646/zootaxa.3857.2.1, http://zenodo.org/record/4929981, {"references":["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.","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.","Jullien, J. & Calvet, L. (1903) Bryozoaires provenant des campagnes de l'Hirondelle (1886 - 1888). Resultats des campagnes scientifiques accomplies sur son yacht par Albert 1 er, Prince Souverain de Monaco, 1903, 23, 1 - 188.","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. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1909. tb 00458. x","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] http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1929. tb 01453. x","Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14 - 17.","Soule, D. F. & Soule, J. D. (1975) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Ortmann, A. (1890) Die Japanische Bryozoenfauna. Bericht uber die von Herrn Dr. L. Doderlein im Jahre 1880 - 81 gemachten Sammlungen. Archiv fur Naturgeschichte, 54, 1 - 74.","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. http: // dx. doi. org / 10.1080 / 00222938609460142","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.","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. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1927. tb 00381. x","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.","Manzoni, A. (1875) I briozoi del pliocene antico di Castrocaro. Gamberini & Parmeggiani, Bologna, 64 pp."]}

Mead, A., Carlton, J.T., Griffiths, C.L., Rius, M. (2011): Introduced and cryptogenic marine and estuarine species of South Africa. Journal of Natural History 45 (39-40): 2463-2524, DOI: 10.1080/00222933.2011.595836, URL: http://dx.doi.org/10.1080/00222933.2011.595836

Lepraliomorph-grade ascophorans are the dominant group of bryozoans in most modern faunas. Here we describe five new species (Parasmittina alba sp. nov., Microporella proxima sp. nov., Reteporella antennata sp. nov., Stephanollona robustaspinosa sp. nov. and Turritigera buski sp. nov.) and record an additional five lepraliomorph species [Watersipora subovoidea (d'Orbigny, 1852), Schizoporella errata (Waters, 1878), Arthropoma cecilii (Audouin, 1826), Celleporina diota (Marcus, 1938) and Reteporellina evelinae Marcus, 1955] from the coastal waters off Rio de Janeiro State in Brazil. This increases the biodiversity of the bryozoan fauna from Brazil, and the number of species known from Rio de Janeiro State now totals 79.

Watersipora subtorquata (d���Orbigny, 1852) (Figs 3, 4 A, C, E, F) Escharina torquata [Milne] Edwards: d���Orbigny 1842, pl. 4, fig. 3; 1847: 11. Schizoporella atrofusca Busk: Hincks 1868: 269, pl. 10, figs 4���5. Schizoporella aterrima Ortmann, 1890: 49, pl. 3, fig. 36. Lepralia ? cucullata: Waters 1909: 150, pl. 15, figs 1, 5 [pars]. Watersipora cucullata: Hastings 1930: 729 [pars], pl. 15, figs 102���104 only. Watersipora cucullata: Marcus 1937: 118, pl. 24, fig. 63 A, B. Watersipora cucullata: Mawatari 1952: 14, fig. 1 A���E (not 1 F). Watersipora cucullata var. watersi Mawatari, 1952: 16, fig. 1 G [?] Dakaria subovoidea: Harmer 1957: 1022, pl. 69, figs 11���13. Watersipora subtorquata: Ryland 1974: 345, fig. 3 C. ��� Watersipora subovoidea ��� fide Harmer: Soule & Soule 1976: 302, pl. 3, fig. 4; pl. 4, fig. 4. Watersipora subtorquata: Soule & Soule 1976: 302, pl. 2, fig. 3; pl. 3, fig. 3; Gordon 1989: 40, pl. 20, B���H); Seo 1999: 222, fig. 1; Taylor & Gordon 2002: 4, reproducing d���Orbigny���s original figures (fig. 1 A, B) and giving an SEM of the holotype (Fig. 1 C); Florence et al. 2007: 39, fig. 14 I J. Material examined. All specimens are in the Natural History Museum, London. Low Isles, Queensland, Australia (Great Barrier Reef), July 1972, JSR (2 slides: 1 intact, NHM 2007.12.14.9; 1 cleaned, NHM 2007.12.14.10) (Ryland 1974); Bay of Arcachon, on shell (NHM 2007.12.14.11), HDeB, August 2003 (1 slide, decalcified, NHM 2007.12.14.12); St-Jacut-de-la-Mer, Brittany, France, HDeB, April 2005 (on oyster shell, NHM 2007.12.14.13; SEMs; 2 slides: 1 cleaned, NHM 2007.12.12.14; 1 decalcified, NHM 2007.12.14.15); St Peter Port harbour, Guernsey, Channel Islands, May 2007, RL (SEM; 3 slides: 1 intact, NHM 2007.12.14.16; 1 cleaned, NHM 2007.1.14.17; 1 decalcified, NHM 2007.12.14.18). Description. Colonies encrusting, subcircular or with broad, spreading lobes, often overgrowing older layers; or erect with bilaminar lobes; sometimes with a broad and extensive orange-red cortical zone, paling and becoming greyish away from the edges; or dark sepia, black or deep purple with a narrow, orange growing margin (colours fade with drying). Zooids subrectangular or slightly hexagonal, sometimes narrower proximally (usually when associated with row division); large, distinct, about twice as long as wide, 750���1100 (mean ~ 950) �� 300���600 (mean ~ 430) ��m (globally), variable within and between colonies. Frontal skeletal wall rather flat, perforated by numerous large round pseudopores, 20���30 ��m diameter; covered by a rather transparent layer with a shiny cuticle. Orifice large, oval, somewhat wider than long; 130���260 (mean ~ 230) ��m; occupying> 10 % of the total zooid area; with a proximal sinus demarcated by condyles, ~ 55 �� 115 ��m; surrounding rim variably developed, sometimes in the form of paired mucrones beside the sinus, often crestlike distally; a pair of areolae proximolateral to the orifice, one each side, about the size of a pseudopore but containing a multiporous septulum. Condyles in the form of narrow shoulders distal to and slightly deeper than the angle of the orifice rim that separates anter from sinus. Operculum strongly pigmented, with a dark broad, biconcave band proximally, gradually spreading around paired clear areas. Polypides with orange lophophores 740���850 ��m long, ~ 24 tentacles (Gordon 1989). Variation and remarks. The operculum, with the dark biconcave central band that spreads distally around a pair of subcircular lateral ���windows���, is diagnostic but the details cannot be seen clearly without special preparation. In the orange colonies from Guernsey the dark band is narrow, about the width of the sinus at the waist (Fig. 4 F); in the black colonies from St-Jacut and the Bay of Arcachon the dark band is noticeably broader, greater than the width of the sinus, and its lateral margins are more sharply defined (Fig. 4 E). Hincks (1886) named a variety labiosa from the Arabian Sea, with the orificial rim raised as mucrones on either side of the sinus, which appears referable to this species and the counterpart of W. subovoidea var. labiosa Calvet (see above). Ortmann���s (1890) Schizoporella aterrima, clearly belonging to Watersipora, is unrecognizable to species. It was included by Mawatari (1952) in the synonymy of W. cucullata but his account is clearly based on W. subtorquata (distinguished as var. watersi, see above) as well as W. subovoidea (as W. cucullata). The illustrated Japanese specimens are W. subtorquata and it is unclear whether any of his local material was W. cucullata., Published as part of Ryland, John S., Blauwe, Hans De, Lord, Richard & Mackie, Joshua A., 2009, Recent discoveries of alien Watersipora (Bryozoa) in Western Europe, with redescriptions of species, pp. 43-59 in Zootaxa 2093 on pages 55-56, DOI: 10.5281/zenodo.274831, {"references":["Hincks, T. (1868) The Polyzoa of the Adriatic: a supplement to Prof. Heller's ' Die Bryozoen des adriatischen Meeres', 1867. Annals and Magazine of Natural History, ser. 5, 17, 254 - 271.","Ortmann, A. (1890) Die Japanische Bryozoenfauna. Bericht uber die von Herrn Dr. L. Doderlein im Jahre 1880 - 81 gemachten Sammlungen. Archiv fur Naturgeschichte, 54, 1 - 74.","Waters, A. W. (1909) Cheilostomata. Reports on the marine biology of the Sudanese Red Sea. XII. The Bryozoa. Part I. Cheilostomata. Journal of the Linnean Society (Zoology), 31, 123 - 181.","Hastings, A. B. (1930) Cheilostomatous Polyzoa from the vicinity of the Panama Canal collected by Dr. C. Crossland on the cruise of the S. Y. ' St. George'. Proceedings of the Zoological Society of London, 1929, 697 - 740.","Marcus, E. (1937). Bryozoarios Marinhos Brasileiros I. Boletim da Faculdada de Filosofia Ciencias e Letras Universidada de Sao Paulo, Serie Zoologica, 1, 5 - 224.","Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14 - 17.","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition. Part 4. Cheilostomata Ascophora II. Siboga Expeditie, 28 D, 641 - 1147.","Ryland, J. S. (1974) Bryozoa in the Great Barrier Reef province. In: Proceedings of the Second International Reef Symposium Volume 2. Cameron, A. M. et al. Brisbane, The Great Barrier Reef Committee. Pp. 341 - 348.","Soule, D. F. & Soule, J. D. (1976) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","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.","Seo, J. E. (1999) Taxonomic review of Korean Watersipora (Bryozoa, Gymnolaemata, Cheilostomata). Korean Journal of Systematic Zoology 15, 221 - 229.","Taylor, P. D. & Gordon, D. P. (2002) Alcide d'Orbigny's work on Recent and fossil bryozoans. Comptes Rendus Palevol, 1, 533 - 547.","Florence, W. K., Hayward, P. J. & Gibbons, M. J. (2007) Taxonomy of shallow-water Bryozoa from the westcoast of Africa. African Natural History, 3, 1 - 58."]}

Watersipora subovoidea (d���Orbigny, 1852) (Fig. 4 C, D, G, H) Cellepora subovoidea d���Orbigny, 1852: 402. Lepralia cucullata Busk, 1854: 81, pl. 96, figs 4, 5. Schizoporella cucullata var. labiosa Calvet, 1903: 141 ���142, pl. 16, fig. 7 a���c. Smittia (Watersipora) cucullata: Neviani 1895: 83. Lepralia ? cucullata: Waters 1909: 150, pl. 15, figs 2���4 [pars]. Watersipora cucullata: Hastings 1930: 729 [pars], pl. 15, figs 97���101 (not 102���104); Soule & Soule 1976: 306, pl. 2, figs 1���2; pl. 3, fig. 1; pl. 4, fig. 3; also pl. 3, fig. 2, as Cellepora ovoidea, from Savigny���s (1817) plate. [? Not] Dakaria subovoidea: Harmer 1957: 1022, pl. 69, figs 11, 12, 14. Watersipora subovoidea: Gautier 1962: 183; Ryland 1965: 68, figs 33 c, d; 1974: 345, fig. 3 A; Soule & Soule 1976: 302, pl. 3, fig. 4; pl. 4, fig. 4; Zabala 1986: 396, fig. 129; Hayward & McKinney 2002: 63, fig. 29 A, B. Material examined. All specimens are in the Natural History Museum, London. Naples, Italy, 20 - 10-1960, JSR (1 slide, intact, NHM 2007.12.14.2); Arrawarra, New South Wales, Australia, January 1772, JSR (3 slides: 1 cleaned, NHM 2007.12.14.3; 2 decalcified, NHM 2007.12.14.4��� 5) (Ryland 1974); Port of Genoa, Italy, 12 - 11-2007, M. Faimali (3 slides: 1 cleaned, NHM 2007.12.14.6; 2 decalcified, NHM 2007.12.14.7��� 8). Description. Colonies encrusting, unilaminar, multilaminar or sometimes foliaceous, brownish-purple to almost black in life (fading to grey when dried), with the growing margin orange. Zooids subrectangular or slightly hexagonal, sometimes narrower proximally (usually when associated with row division); large, distinct, about twice as long as wide, 700���1100 (mean ~ 900) �� 350���650 (mean ~ 450) ��m (globally; Table 2), variable within and between colonies. Frontal skeletal wall rather flat, perforated by numerous large round pseudopores 20���30 ��m diameter; in some zooids two or more distal pores, largely concealed by the orificial rim, may be areolar septular pores; covered by a variably pigmented layer and a shiny, transparent cuticle. Orifice large, subcircular to oval, slightly wider than long; 200���250 (mean ~ 205) �� 190���250 (mean ~ 230) ��m; occupying Variation and remarks. The rim of the orifice may be somewhat elevated on either side of the sinus, as in the form distinguished as var. labiosa by Calvet (1903, pp. 141, 142), from the intertidal of Graciosa, Azores. Waters (1909) confounded two species under his Lepralia cucullata ― his Naples specimens were correctly identified and are now therefore referred to W. subovoidea, but those from the Red Sea (Plate 15, figs 1, 5), later distinguished as var. watersi by Mawatari (1952), are clearly referable on the basis of the operculum to W. subtorquata., Published as part of Ryland, John S., Blauwe, Hans De, Lord, Richard & Mackie, Joshua A., 2009, Recent discoveries of alien Watersipora (Bryozoa) in Western Europe, with redescriptions of species, pp. 43-59 in Zootaxa 2093 on page 54, DOI: 10.5281/zenodo.274831, {"references":["Calvet, L. (1903) Deuxieme Partie. In: Jullien, J. & Calvet, L. Bryozoaires provenant des campagnes de l' Hirondelle (1886 - 1888). Resultats des campagnes scientifiques accomplies sur son yacht par Albert 1 er, Prince Souverain de Monaco, 1903, 23, 1 - 188.","Neviani, A. (1895) Briozoi fossili della Farnesina e Monte Mario presso Roma. Palaeontographia italica, 1, 77 - 140.","Waters, A. W. (1909) Cheilostomata. Reports on the marine biology of the Sudanese Red Sea. XII. The Bryozoa. Part I. Cheilostomata. Journal of the Linnean Society (Zoology), 31, 123 - 181.","Hastings, A. B. (1930) Cheilostomatous Polyzoa from the vicinity of the Panama Canal collected by Dr. C. Crossland on the cruise of the S. Y. ' St. George'. Proceedings of the Zoological Society of London, 1929, 697 - 740.","Soule, D. F. & Soule, J. D. (1976) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition. Part 4. Cheilostomata Ascophora II. Siboga Expeditie, 28 D, 641 - 1147.","Gautier, Y. V. (1962) Recherches ecologiques sur les Bryozoaires chilostomes en Mediterranee occidentale. Receuil des Travaux de la Station marine d'Endoume, Fasc. 38, Bull. 24, 1 - 434.","Ryland, J. S. (1965) Polyzoa. OECD Catalogue of Main Marine Fouling Organisms, 2, 1 - 83.","Zabala, M. (1986). Fauna dels Briozous dels Paisos Catalans. Barcelona, Institut d'Estudis Catalans. 833 pp.","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.","Ryland, J. S. (1974) Bryozoa in the Great Barrier Reef province. In: Proceedings of the Second International Reef Symposium Volume 2. Cameron, A. M. et al. Brisbane, The Great Barrier Reef Committee. Pp. 341 - 348.","Mawatari, S. (1952) On Watersipora cucullata (Busk). I. Systematic study. Miscellaneous Reports of the Research Institute of Natural Resources (Tokyo), 25, 14 - 17."]}

Genus Watersipora Neviani, 1895 (Cheilostomatida: Smittoidea: Watersiporidae) Diagnosis. Colonies encrusting, uni- to multilaminar, or suberect to erect, bilaminar; autozooids with an evenly perforated, often transparent, frontal shield with the compensation sac (ascus) below it (Hyman 1959, figure 120 D, E); a pair of small areolar septula, each multiporous, positioned beside the orifice, also appear to be a generic character; vertical walls perforated by large, multiporous rosette plates (septula). Primary orifice often sinusoid, with prominent condyles (cardellae); no spines. Opercula with straight or curving, lengthwise sclerites, often darkly pigmented (especially between the sclerites); sometimes with a pair of internal proximobasal knobs showing from above as translucent spots (lucidae). No avicularia. No external ovicells; each embryo brooded in a distally situated internal embryo sac (Waters 1909, pl. 15, 4). Ancestrula oval in outline, with the form of later zooids. Type species. Lepralia cucullata Busk, 1854. 2.0 1.9 R 2 K 35 ) R 3 2 S 17 1.8 1000 M 25 K 9 K 44 S 5 (area 1.7 M 15 S M 1 35 K 3 S 7 orifice 1.6 M 68 K 18 S 11 S 3 M 5 S 8 Log S 6 M 65 S 12 R 1 S 10 S 4 S 9 S 2 M 70 S 14 1.5 R 4 S 13 S 15 M 55 S 16 1.4 S 18 Remarks: There is a long-standing problem with the usage of the name W. subovoidea. This problem stems from Harmer (1957), who observed that ��� Cellepora ovoidea Audouin, 1826 [renamed Cellepora subovoidea by d���Orbigny (1852) because C. ovoidea was a preoccupied name], seems to be a synonym of the species commonly described as Lepralia cucullata [Busk, 1854]���. Evidently applying the Principle of Priority (International Commission on Zoological Nomenclature [ICZN] 1999), Harmer reverted to the earlier name. This was unfortunate for, although workers had muddled the identity of L. cucullata, Busk���s species could be clearly established by his Aegean Sea type specimen, whereas Audouin���s species is unrecognizable. It was described by Audouin from a picture by Savigny (1817, pl. 8, fig. 1, no text), which has subsequently been reproduced by Soule (1976) and more recently by d���Hondt (2006). It may reasonably be assumed to represent a Watersipora but is too stylized to allow association with any particular species. Its undisclosed locality is presumably the Egyptian coast of either the eastern Mediterranean or the Red Sea (then, of course, not connected by the Suez Canal). The identity of Busk���s L. cucullata is, as explained, clear, following the account of Hastings (1930), irrespective of possible misidentifications in other parts of the world. The species is the one common throughout the entire Mediterranean basin (Gautier 1962, unillustrated; Ryland 1965, with line drawings; Zabala 1986, including Ryland���s fig. 33 d; Zabala & Maluquer 1988, unillustrated; Hayward & McKinney 2002, with SEMs). These authors have accepted Harmer���s (1957) synonymy and have used W. subovoidea in the sense of W. cucullata (Busk). In the interests of nomenclatural stability we believe that the best solution is to designate Busk���s Aegean Sea specimen, NHM 1854.11. 15.189, as the neotype of W. subovoidea (d���Orbigny) in accordance with Article 75.3 of the International Code (ICZN 1999). W. cucullata and W. subovoidea thus become objective synonyms and priority dictates that the latter should be the name used. Existing usage of the name subovoidea is thereby stabilized and the designation ��� W. subovoidea sensu Ryland (1974) ��� becomes un-necessary. Lepralia complanata Norman (1864), currently placed in Watersipora (Hayward & Ryland 1999), with a campanulate orifice, does not appear obviously congeneric with the arcuata / subovoidea / subtorquata group, to which the type species belongs., Published as part of Ryland, John S., Blauwe, Hans De, Lord, Richard & Mackie, Joshua A., 2009, Recent discoveries of alien Watersipora (Bryozoa) in Western Europe, with redescriptions of species, pp. 43-59 in Zootaxa 2093 on pages 52-54, DOI: 10.5281/zenodo.274831, {"references":["Neviani, A. (1895) Briozoi fossili della Farnesina e Monte Mario presso Roma. Palaeontographia italica, 1, 77 - 140.","Hyman, L. H. (1959) The invertebrates: smaller coelomate groups. New York, McGraw-Hill. viii + 783 pp.","Waters, A. W. (1909) Cheilostomata. Reports on the marine biology of the Sudanese Red Sea. XII. The Bryozoa. Part I. Cheilostomata. Journal of the Linnean Society (Zoology), 31, 123 - 181.","Harmer, S. F. (1957) The Polyzoa of the Siboga Expedition. Part 4. Cheilostomata Ascophora II. Siboga Expeditie, 28 D, 641 - 1147.","Savigny, J. - C. (1817). Description de l'Egypte ou recueil des observations et des recherches qui ont ete faites en Egypte pendant l'expedition de l'Armee Francaise. Atlas, Histoire naturelle (1809 - 17). Paris.","Soule, D. F. & Soule, J. D. (1976) Species groups in Watersiporidae. Bryozoa 1974. Documents des Laboratoires de Geologie de la Faculte des Science de Lyon, h. s. 3, 2, 299 - 309.","Hastings, A. B. (1930) Cheilostomatous Polyzoa from the vicinity of the Panama Canal collected by Dr. C. Crossland on the cruise of the S. Y. ' St. George'. Proceedings of the Zoological Society of London, 1929, 697 - 740.","Gautier, Y. V. (1962) Recherches ecologiques sur les Bryozoaires chilostomes en Mediterranee occidentale. Receuil des Travaux de la Station marine d'Endoume, Fasc. 38, Bull. 24, 1 - 434.","Ryland, J. S. (1965) Polyzoa. OECD Catalogue of Main Marine Fouling Organisms, 2, 1 - 83.","Zabala, M. (1986). Fauna dels Briozous dels Paisos Catalans. Barcelona, Institut d'Estudis Catalans. 833 pp.","Zabala, M. & Maluquer, P. (1988) Illustrated keys for the classification of Mediterranean Bryozoa. Treballs del Museu de Zoologia (Barcelona), 1988, 1 - 294.","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.","Ryland, J. S. (1974) Bryozoa in the Great Barrier Reef province. In: Proceedings of the Second International Reef Symposium Volume 2. Cameron, A. M. et al. Brisbane, The Great Barrier Reef Committee. Pp. 341 - 348.","Hayward, P. J. & Ryland, J. S. (1999) Cheilostomatous Bryozoa. Part 2, Hippothooidea - Celleporoidea. Synopses of the British Fauna, n. s., 14, 1 - 416."]}

Dick, Matthew H., Tilbrook, Kevin J., Mawatari, Shunsuke F. (2006): Diversity and taxonomy of rocky-intertidal Bryozoa on the Island of Hawaii, USA. Journal of Natural History 40 (38-40): 2197-2257, DOI: 10.1080/00222930601062771, URL: http://dx.doi.org/10.1080/00222930601062771