taxonID	type	description	language	source
6E0B87BA463B4C3614B5EB7AFAE3FD53.taxon	description	Critical to understanding the correct name of the common fouling species of Watersipora is establishing the identity of W. subtorquata (d’Orbigny, 1852). Specimens of a bryozoan from Rio de Janeiro described by d’Orbigny (1842) as Escharina torquata (Lamouroux, 1825) were later renamed by him (d’Orbigny 1852, p. 399) as Cellepora subtorquata, presumably to avoid homonymy with another species he referred to as Cellepora torquata (Quoy & Gaimard, 1827) (see d’Orbigny 1852, p. 403). Waters (1879) synonymised d’Orbigny’s Cellepora subtorquata with Lepralia cucullata Busk, 1854 but inexplicably applied the name of the junior synonym L. cucullata for his specimens from Naples. The first revision of d’Orbigny’s type was undertaken subsequently by Waters (1905) who studied d’Orbigny’s material deposited at the MNHN. He noted the high quality of d’Orbigny’s original figures but remarked that, in the original specimens, the projecting lateral wings on each side of the sinus were more developed than those depicted by d’Orbigny (1842). Taylor & Gordon (2002) reviewed the bryozoan work of Alcide d’Orbigny (1802 – 1857), publishing a scanning electron micrograph of d’Orbigny’s type specimen (Taylor & Gordon 2002, fig. 1 C) in which it is possible to recognize some of the diagnostic features (orifice shape and frontal pseudopores) depicted in the original figures of this species. The type specimen of d’Orbigny (MNHN, d’Orbigny Collection 13637) (Figs 1 – 5) comprises a large colony with some intact opercula (Figs 1 – 2); the projecting proximolateral wings are well developed in most zooids (Fig. 4), as observed by Waters (1905), but with increasing calcification can become obscured, shallow and inconspicuous in frontal views of the orifice (Figs 3, 5). The subcircular orifice (Fig. 5) was well figured in d’Orbigny’s plates; the orifice is slightly wider than long, with a U-shaped sinus demarcated by triangular condyles that project distomedially. Comparisons of d’Orbigny’s type specimen and the specimens identified by Marcus (1937, 1938) from Brazil under the name Watersipora cucullata (NHMUK 1948.2.16.18, and uncatalogued specimens deposited at MZUSP) indicate that they represent the same species, as has been suggested by Gordon (1989) and Ryland et al. (2009). Marcus (1937, pl. 24, fig. 63 A, B) showed the orifice to have well-developed proximolateral wings in Brazilian colonies. In addition, the operculum in the Brazilian specimens is characterized by a parallel-sided dark band of two proximal lucidae adjacent to the condyles (Marcus 1937, pl. 24, fig. 63 A, B). The same operculum shape and lucidae were observed in Rio de Janeiro specimens figured by Ramalho et al. (2009, fig. 3 D) in material they named Watersipora subovoidea. Ramalho et al. (2011) followed Ryland et al. (2009) in using the name W. subovoidea for specimens with “ triangular, tooth-like condyles, located distomedially [sic], and a strongly pigmented operculum with a parallel-sided dark central band ”. All characteristics described for W. subovoidea by Ryland et al. (2009) suggest that their specimens actually belong to W. subtorquata, and restudy of the material they analyzed (NHMUK 2007.12.14.2 – 8) confirms this supposition. However, neither figures nor descriptions of Busk’s type specimen of Lepralia cucullata (NHMUK 1854.11.15.189), designated by Ryland et al. (2009: 54) as the neotype of Watersipora subovoidea, were given by Ryland et al. (2009), a situation which we rectify below.	en	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
6E0B87BA46384C3714B5ED62FB50FE24.taxon	description	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.	en	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
6E0B87BA46394C3714B5EC4AFBCAF864.taxon	type_taxon	Type species. Lepralia cucullata Busk, 1854, by original designation.	en	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
6E0B87BA46394C3714B5EC4AFBCAF864.taxon	diagnosis	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.	en	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
6E0B87BA46394C3714B5EC4AFBCAF864.taxon	discussion	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, 1969 a, 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, 1969 b has dimorphic ovicellate zooids (see Banta 1969 b, figs 2 – 4).	en	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
6E0B87BA463E4C3F14B5EEB1FE95FB41.taxon	description	(Figs 1 – 5, 12 – 16, 18 – 24, 67, 70; Table 1)	en	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
6E0B87BA463E4C3F14B5EEB1FE95FB41.taxon	materials_examined	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 34 a, 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, F 78, O’Donoghue coll. NHMUK 1963.8.2.41 pt, dry, Watersipora subovoidea, J. Soule & D. Soule det., Alexandria, Egypt, Station 34 a, Eastern Harbour, O’Donoghue coll. NHMUK 1973.1. 10.3, dry, Accra, Ghana, No. 13 B, 14 February 1949, Bassindale coll. NHMUK 1973.1.10.4 pt, 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. 2 a, 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, 23 m, 15 September 1978, Dumont coll. NHMUK 2012.6. 30.16, dry, Watersipora subovoidea, Shaab Baraja, Sudan, Red Sea, 25 º 52 ’ N, 37 º 24 ’ 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. 13 B, 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.	en	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
6E0B87BA463E4C3F14B5EEB1FE95FB41.taxon	description	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.	en	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
6E0B87BA463E4C3F14B5EEB1FE95FB41.taxon	discussion	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. 1 G) 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. 156 B) 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).	en	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
6E0B87BA463E4C3F14B5EEB1FE95FB41.taxon	distribution	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).	en	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
6E0B87BA46314C3B14B5EB12FC78F900.taxon	description	(Figs 6 – 9, 25 – 34, 65; Table 2)	en	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
6E0B87BA46314C3B14B5EB12FC78F900.taxon	materials_examined	Material examined. Lectotype (chosen by Soule & Soule 1975): NHMUK 1854.11. 15.189 (specimen mounted on one dry slide and two balsam slides), Lepralia cucullata, 1854, G. Busk det., Aegean Sea, E. Forbes. Paralectotypes: NHMUK 1899.7. 1.1398, dry, same data as for lectotype; NHMUK 2012.6. 30.1, dry slide, same data as for lectotype. Other material: NHMUK 1899.5. 1.456, Schizoporella atrofusca, dry slide, T. Hincks det., Adriatic, Pieper coll. NHMUK 1899.5. 1.975, dry slide, Schizoporella atrofusca, T. Hincks det., Adriatic. NHMUK 1965.8. 14.10, dry slide, Watersipora cucullata, Balearic Islands, Mediterranean, Cox coll.	en	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
6E0B87BA46314C3B14B5EB12FC78F900.taxon	description	Description. Colony encrusting, multiserial, unilamellar; blackish in colour. Zooids subrectangular to hexagonal, separated by slightly raised grooves. Frontal shield slightly more convex distally than proximally, with small tubercles and uniformly perforated by round pseudopores about 20 µm diameter; two latero-oral intrazooidal septula, small, near the lateral zooidal margin proximolateral to the orifice; each with 2 – 4 small pores; frontal shield obscured by thick, black epitheca. Orifice large, slightly wider than long, with a well-defined, broad, shallow, U-shaped sinus demarcated by triangular projections; orificial rim robust and thickened around whole orifice, sometimes better developed proximally than distally; condyles upturned, large and conspicuous. Operculum with broad, parallel-sided dark central band and slightly thinner lateral area. Avicularia absent. Ooecia absent.	en	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
6E0B87BA46314C3B14B5EB12FC78F900.taxon	discussion	Remarks. Watersipora subovoidea has been reported in subtropical and tropical waters, i. e. Brazil, Florida and Australia (Mackie et al. 2006; Geller et al. 2008; Ryland et al. 2009), but the great majority of these records belong to W. subtorquata (see above). D’Orbigny (1852) did not give any description or figures for Cellepora subovoidea, and according to the ICZN Article 75.3.6 “ evidence that the neotype came as nearly as practicable from the same original type locality ” should be provided, which is not apparent in the Ryland et al. (2009) paper. Thus, we suggest setting aside the neotype selection of Cellepora subovoidea (NHMUK 1854.11.15.189) made by Ryland et al. (2009). Watersipora cucullata is characterized by a suborbicular orifice with a shallow, wide sinus demarcated by triangular projections (Figs 8, 27, 29, 31, 32, 34), large, conspicuous condyles (Figs 8, 27, 29, 31), and a small latero-oral intrazooidal septula on each side of the zooid (Fig 8, 29, 34). Latero-oral intrazooidal septula are also present in Watersipora aterrima and W. subatra, but these species are distinguished by the shape of the orifice and condyles (see below). Watersipora cucullata differs from W. nigra in having smaller zooids and orificial area (see Canu & Bassler, 1930, p. 26; Fig. 11; Table 2) and paired intrazooidal septula, near the lateral zooidal margin, proximolateral to the orifice. Watersipora atrofusca (Busk, 1856) (Fig. 10; Table 2), known from Mazatlan (Mexico), resembles W. cucullata in colony shape but differs in having a smaller, almost circular orifice and in lacking intrazooidal septula.	en	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
6E0B87BA46314C3B14B5EB12FC78F900.taxon	distribution	Distribution. Mediterranean (including the Adriatic and Aegean seas).	en	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
6E0B87BA46354C2014B5E8D6FE2DF9D6.taxon	description	(Figs 39 – 53, 66, 69; Table 3)	en	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
6E0B87BA46354C2014B5E8D6FE2DF9D6.taxon	materials_examined	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.	en	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
6E0B87BA46354C2014B5E8D6FE2DF9D6.taxon	description	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.	en	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
6E0B87BA46354C2014B5E8D6FE2DF9D6.taxon	discussion	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. 3 A, 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.	en	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
6E0B87BA46354C2014B5E8D6FE2DF9D6.taxon	distribution	Distribution. NE Atlantic (Ireland and British Isles), Indo-West Pacific (Indonesia) and Pacific (New Zealand, Australia and California).	en	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
6E0B87BA462E4C2314B5E8E1FE35FB3C.taxon	description	(Figs 54 – 58; Table 4)	en	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
6E0B87BA462E4C2314B5E8E1FE35FB3C.taxon	materials_examined	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 – 50 m, dredged, 22 October 1996, T. Kato coll.	en	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
6E0B87BA462E4C2314B5E8E1FE35FB3C.taxon	etymology	Etymology. Named for Japanese bryozoologist Prof. Shunsuke F. Mawatari.	en	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
6E0B87BA462E4C2314B5E8E1FE35FB3C.taxon	description	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.	en	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
6E0B87BA462E4C2314B5E8E1FE35FB3C.taxon	discussion	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.).	en	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
6E0B87BA462E4C2314B5E8E1FE35FB3C.taxon	distribution	Distribution. Japan.	en	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
6E0B87BA462D4C2E14B5EACEFEC5FDBB.taxon	materials_examined	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 – 41 pt., 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 1921.5.24.18 C, 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 1981.4.1.3 pt, dry and balsam slide, Watersipora subovoidea, Ceylon [Sri Lanka], L. R. Thornely, Rep. Pearl Oyster Fish., 26, p 120. NHMUK 2012.6. 30.1 [part of NHMUK 1912.12.21.1019], Watersipora cucullata, Naples, Italy, A. M. Norman.	en	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
6E0B87BA462D4C2E14B5EACEFEC5FDBB.taxon	description	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.	en	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
6E0B87BA462D4C2E14B5EACEFEC5FDBB.taxon	etymology	Etymology. Named for John D. Soule (1920 – 2001) and Dorothy F. Soule (1923 – 2005), in recognition of their contributions to bryozoan taxonomy.	en	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
6E0B87BA462D4C2E14B5EACEFEC5FDBB.taxon	discussion	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.	en	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
6E0B87BA462D4C2E14B5EACEFEC5FDBB.taxon	distribution	Distribution. Atlantic (Azores, Cape Verde and Senegal), Mediterranean (Naples) and Indian Ocean (Sri Lanka and India).	en	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
